Written by David R. Curtis and Per Andersen.

Introduction

Sir John Eccles, internationally recognized for his remarkable and outstanding impact on the neurosciences for more than six decades, died on 2 May 1997 at the age of 94. He carried out his research in Oxford, Sydney, Dunedin, Canberra, Chicago and Buffalo from 1927 until 1975 (441). His numerous scientific papers and books, arising from detailed and pioneering experimental studies of synaptic mechanisms and the organization of neurones in the mammalian central nervous system, continue to have a major influence on brain research. Furthermore, his writings on the mind-brain interaction generated wide interest and debate. Eccles also made his mark as an administrator, particularly at the Australian National University and the Australian Academy of Science, of which he was a Foundation Fellow and the second President.

Early years and family background

John Carew Eccles was born on 27 January 1903 at Northcote, a suburb of Melbourne. Both his father, William James Eccles, and his mother, Mary (née Carew) were born in Victoria and were school teachers. The Eccles family came from Lancashire in England, his paternal grandfather Henry Basil Eccles having migrated to Victoria in 1849 and his grandmother Mary Jane Ingram from Limerick in Ireland in the same year. His maternal grandfather John Carew migrated to Victoria from Tipperary in Ireland in 1852, his grandmother Harriet Elizabeth Merry from Berkshire in England in 1858. Eccles's sister, Rosamond, to whom he was closely attached, was born in 1904.

At the age of 12, Eccles began his secondary schooling at Warrnambool High School. After four years, prior to entering the University of Melbourne, he studied science and mathematics for another year at Melbourne High School. He headed the school at the final State-wide examination, shared the State geometry prize and gained a Senior Scholarship to the University. Although deeply interested in mathematics, Eccles chose medicine and commenced his five-year course early in 1920 at the age of 17. He lived at home and attended tutorials at Newman College. His continuing academic successes attracted the attention of a number of professors, including W.A. Osborne (Physiology). As a medical student he was active in university societies and sport, gaining a full blue in athletics for his Australian universities record in pole vaulting.

Eccles also developed his interests in the arts. His reading of Darwin's Origin of Species in the first-year zoology course led to wide reading of classical and contemporary works in philosophy dealing with the mind-brain problem. Unable to obtain a satisfactory explanation of the interaction between the mind and the brain and realizing that so little was known about the brain itself, he decided as a medical student to become a neuroscientist (see 425). After reading CS Sherrington's 1906 book, The Integrative Action of the Nervous System, Eccles resolved to achieve a Rhodes Scholarship to work with him at Oxford University. He later described Sherrington as 'the one man in the world whom I wished to have as my master' (441).

Eccles completed his medical course in February 1925, gaining First Class Honours and First Place, the exhibition in eight of ten subjects and several clinical prizes, and graduated with the degrees of Bachelor of Medicine and Bachelor of Surgery. He had already been awarded the Rhodes Scholarship for Victoria in November 1924. After six months as a Resident Medical Officer at St Vincent's Hospital, he left Melbourne at the end of August 1925 and arrived in Oxford early in October.

Oxford

Eccles had been accepted by Magdalen College, of which Sir Charles Sherrington was a Fellow. A letter of introduction from Sherrington's friend Professor Osborne initiated Eccles's close and deeply influential friendship with Sir Charles, which continued until the latter's death (441,468, 505). Initially, he spent two years studying for the Final Honours School in Physiology and Biochemistry, and over this period at Magdalen he read widely in the neurophysiological, biochemical, philosophical and theological literature. In mid-1927 he was awarded First Class Honours, the Christopher Welch Scholarship and a Gotch Prize. He commenced his DPhil under the supervision of Sherrington in the autumn of 1927 and moved to Exeter College, having been appointed a Junior Research Fellow for five years.

Eccles has vividly described 'life in Sherrington's laboratory' over the period 1925-1935 which was enriched by numerous visitors from Europe and North America (441, Chap.5 of 468, and 505). At this time there was much speculation regarding the nature of the transmission process at central synapses. Strong evidence for chemical transmission at peripheral excitatory and inhibitory synapses had been provided, particularly by the findings of O Loewi and of HH Dale and his colleagues. Nevertheless, central synaptic transmission was widely considered to be similar to the electrical propagation of impulses along nerve fibres, although Sherrington (1925) had suggested that the inhibition of spinal reflexes might be mediated by a chemical agent.

Eccles's introduction to research was with EGT Liddell and D Denny-Brown in a study of the effects of electrical stimulation of the cerebellar cortex on spinal reflexes in the cat (3). Later in 1927 he joined RS Creed in a study of inhibition in the spinal cord (1), and in 1928 investigated crossed extensor reflexes with RA Granit. Reflex responses were measured as muscle contractions by means of an optical isometric myograph, the design of which was improved after Eccles discovered that friction in the bearing distorted the records (4).

In mid-1928 Eccles joined Sherrington in a series of experiments. The first led to the discovery of two distinct populations, based on diameter, of motor fibres in peripheral muscle nerves (7). It was not until later that L Leksell (1945) showed these to be the axons of alpha- and gamma-motoneurones, the latter innervating muscle spindle stretch receptor organs. Sherrington and Eccles (see 14) also examined the time course of the 'central excitatory state' (c.e.s.) which Sherrington had proposed underlay the excitation of motoneurones by afferent volleys to the spinal cord, and also of the active 'central inhibitory state' (c.i.s.) associated with the inhibition of flexor reflexes by volleys in contralateral muscle nerves (17). The latter observations provided no support for the then current theory that the inhibition of reflexes resulted from interference with the access of excitatory impulses to the cord.

These experiments finished early in 1931, and were the last in which Sherrington, then aged 74, took an active part. In 1932 the influential monograph Reflex Activity of the Spinal Cord was published by Creed, Denny-Brown, Eccles, Liddell and Sherrington (21) giving an account of studies carried out in the Sherrington 'School' in Oxford over the previous decade. That year Sherrington and ED Adrian from Cambridge shared the Nobel Prize for Physiology or Medicine for 'their discoveries regarding the functions of the neurone'.

Sherrington's Nobel Lecture, entitled 'Inhibition as a coordinative factor', had a major influence on the future research activities of John Eccles, then aged 29. His own 1963 Nobel Lecture (267) was entitled 'The ionic mechanism of postsynaptic inhibition'.

In 1931 Eccles was appointed to the Staines Medical Fellowship at Exeter College. A study with GL Brown of the acetylcholine-mediated vagal inhibition of the heart apparently reinforced his belief that only an electrical process could account for the much shorter delays and duration of transmission at central synapses. In 1932, Eccles, JZ Young and Granit showed, for the first time, that action potentials were conducted in both directions along earthworm giant nerve fibres (20), an observation which, when later extended by others to the giant axon of the squid, was of fundamental significance to the understanding of impulse conduction along axons.

Eccles returned to the study of synaptic transmission in 1935, selecting the superior cervical sympathetic ganglion as a simpler system than the spinal cord. Although he accepted that acetylcholine (ACh) released by presynaptic impulses was the chemical mediator for slow excitation, the failure of the ACh esterase inhibitor physostigmine to prolong the fast component of ganglionic responses strengthened his doubts that the fast excitatory process could be chemical. Accordingly, he introduced the concept of a rapid 'detonator' response by which the action currents of presynaptic impulses directly excited cells at synaptic regions, causing ganglion cells to discharge (41, 506). This proposal was extended to synaptic excitation of striated muscle, of smooth muscle and in the spinal cord (50). The ensuing controversy between Eccles and adherents of chemical synaptic transmission considerably enlivened many meetings of the Physiological Society (429, 506, see Dale 1954).

Eccles continued demonstrating, tutoring, lecturing and supervising undergraduate and DPhil students, and in 1934 achieved a permanent position in Oxford as a Tutorial Fellow at Magdalen College and as a University Demonstrator. In 1935, however, Sherrington retired and was replaced by J Mellanby. Eccles was disappointed about the new directions research in Oxford would take, and was also concerned about the increasing political uncertainty in Europe. Accordingly, he applied successfully for the Directorship of the Kanematsu Memorial Institute of Pathology at Sydney Hospital, then the largest general hospital in Sydney. He resigned his Oxford appointments and arrived in Sydney with his family in August 1937.

Sydney

The Kanematsu Institute (see Courtice 1985) opened in April 1933, initially as a department of diagnostic pathology for the Sydney Hospital. The part-time Director, Dr WK Inglis, had persuaded the Hospital Board that research should be an integral function of the Institute, and funds were sought from the Nuffield Foundation and the government of New South Wales. The position of Director was advertised in August 1935, and the appointee was expected to devote most of his time to some field of medical research, whilst exercising general supervision of the routine work of the Institute. The Hospital Board, however, retained administrative control of the Institute.

Eccles proposed to continue his research on the central and peripheral nervous systems and, by the end of 1937, he had established research facilities on the top floor of the building with the assistance of a grant from the Australian National Health and Medical Research Council (NHMRC). Throughout his period in Sydney he was very concerned about his academic isolation from physiologists at the Medical School of the University. Although he had been elected a Foundation Fellow of the Royal Australasian College of Physicians in 1938 in recognition of his scientific achievements, Eccles and his colleagues never had any formal association with the University. In 1938, 1939, and jointly with B Katz in 1940, however, he gave a series of lectures to the third-year medical students at the University. Thereafter, and until 1943, lectures, demonstrations and discussions took place for interested students at the Institute.

In 1938 Eccles and WJ O'Connor for the first time recorded end-plate potentials electrically from the surface of striated muscle strips. The belief that these were preceded by muscle action potentials, coupled with the effects of physostigmine and the ACh receptor antagonist curare, led Eccles to deny a role for ACh as the neuromuscular transmitter and to claim support for his detonator theory (59). The subsequent detailed study of neuromuscular transmission with B Katz and SW Kuffler, however, led to Eccles's apparent acceptance that ACh was responsible for end-plate potentials set up by motor nerve impulses (68). An investigation of transmission in the cat stellate ganglion appeared to confirm his earlier proposal of dual fast and slow excitatory processes in ganglia, only the latter being mediated by ACh (69,71).

In accord with a Hospital Board desire for clinically relevant research, Eccles also investigated the atrophy of striated muscle which follows disuse or tenotomy (74). The outbreak of war in Europe in 1939 led to some reduction of the Institute's research, and its continuation was severely curtailed from late 1941 by the war in the Pacific region. Eccles, who had been elected FRS in March 1941, became involved in a number of committees and research projects dealing with the problems of vision (70), hearing, noise and communication in aircraft and tanks. He also actively participated with an Army unit responsible for the supply of blood and serum for the armed forces.

In the meantime, however, Eccles's relations with the Hospital Board became less than harmonious. He believed that the Board and the Hospital's honorary medical staff lacked an understanding of the basic neurophysiological research being carried out and of its long-term relevance to clinical medicine. Early in 1943, without consulting Eccles, the Board proposed to add two floors to the Institute building to accommodate resident medical staff. Since this would prevent any future expansion of the research laboratories, Eccles resigned in October 1943 and accepted appointment to the Chair of Physiology in the Medical School of the University of Otago, New Zealand.

Dunedin

Eccles arrived in Dunedin with his family in January 1944. The university, the first in New Zealand, was founded by the Presbyterian Church in 1869 when Dunedin was New Zealand's principal city. The Medical School, the only one in the country, was established in 1876. The Professor of Physiology was also responsible for teaching biochemistry, and Eccles appointed N Edson as Senior Lecturer in Biochemistry. The teaching load was heavy, and considerable changes were made to provide students, over two pre-clinical years, with a scientific basis for the practice of medicine similar to the Final Honours School in Oxford. Eccles also introduced in 1945 a BMedSci degree whereby at the end of the second year a number of the best students spent twelve months on a research project before returning to their medical studies.

A research laboratory was set up with financial support from the Medical Research Council of New Zealand, and Eccles commenced experimentation late in 1944. He recorded ventral root responses to dorsal root stimulation (76) and, with J.L.Malcolm, potentials and 'reflexes' from dorsal roots (77) which had been described earlier by Barron and Matthews (1938). All results were considered to be consistent with an electrical hypothesis of synaptic and neuromuscular transmission that Eccles published in Nature in December 1945 (75), and presented in February 1946 at the New York Academy of Sciences during his first visit to the United States.

Eccles's formal enunciation of his electrical hypothesis was primarily the consequence of his meeting in May 1945 with KR Popper (see Miller 1997). Popper had been a member of the staff of Canterbury University College in Christchurch, New Zealand, since 1937. At the invitation of Eccles and Edson, he spent a week in Dunedin lecturing on, and discussing his views about, the philosophy of science. Eccles was deeply impressed by Popper's main tenet, that scientific hypotheses should be both clearly formulated and testable by experiment, and that the strength of a hypothesis depended on the failure of rigorous investigation to falsify it rather than on evidence which apparently supported it. This meeting, and subsequent meetings in Christchurch, not only led to Eccles's continuing friendship with Popper (see 450, 515, 516), but additionally had a marked impact upon his future research (see 359). To quote Miller, when referring to Popper's lecture course in Dunedin: 'It had the notable effect also of converting a naive believer in induction (as Eccles described himself [see 515]) into one of the most vigorous of all scientific advocates of the method of conjectures and refutations'.

At the time, Eccles considered chemical transmission to play a subordinate or negligible role in sympathetic ganglia, at the neuromuscular junction and in the spinal cord, despite the strong evidence, particularly from the investigations of Dale and his colleagues, that ACh mediated both ganglionic and neuromuscular transmission. Eccles's experimental evidence that ACh was unlikely to be a transmitter in the spinal cord (80) reinforced his opinion that central synaptic transmission was an electrical process. Accordingly, influenced and encouraged by Popper, he stated his hypothesis of electrical excitatory transmission at central and ganglionic synapses, and the neuromuscular junction, in precise terms, and proposed a number of crucial physiological and pharmacological tests (75).

In essence, Eccles replaced his earlier 'detonator' theory with the proposition that presynaptic action currents initiated depolarizing local responses at specialized regions of the postsynaptic membrane. Above a critical level of depolarisation action potentials would be generated. The synaptic delay and the time course of the synaptic potential were accounted for in terms of the time course of the presynaptic action currents and the electrical properties of the postsynaptic membrane. The terminal regions of presynaptic fibres were also proposed to be similarly specialized, so explaining dorsal root potentials (DRPs) and reflexes (DRRs). He also provided explanations for some of the difficulties his hypothesis faced.

Early in 1946 Eccles and C.McC.Brooks tested this hypothesis by recording the electrical events associated with monosynaptic excitation of cat spinal motoneurones. Enamel-insulated metal electrodes were used to record extracellular synaptic and action potentials (focal potentials) generated by presynaptic impulses. The results were interpreted as 'agreeing closely with the predictions of the electrical hypothesis of synaptic transmission' (82, 86). The following year, Brooks and Eccles published in Nature (81) an electrical hypothesis of central inhibition, developed to account for the 'direct' inhibition of spinal reflexes by impulses from antagonistic muscles. Earlier, both B.Renshaw and DPC Lloyd at the Rockefeller Institute had suggested that direct inhibition had the same central latency as that of the monosynaptic excitation of motoneurones.

Eccles's electrical hypothesis proposed that the inhibitory pathway included a short-axon interneurone (Golgi cell) that synapsed upon motoneurones but did not discharge an impulse when excited by inhibitory afferent volleys. Synaptic potentials caused currents to flow through Golgi cell axon terminals and to passively depress motoneurone excitability. This hypothesis was tested by recording synaptic potentials near motoneurones and ventral root reflexes. The results, together with the interaction between synaptic inhibition (88) and antidromic invasion of motoneurones (89), were regarded as being consistent with the Golgi cell hypothesis.

In 1949 Eccles reviewed and restated his electrical hypotheses of synaptic excitation and inhibition in the spinal cord in a slightly modified form (92). His study with W.V.Macfarlane in 1948 of the effects of a number of ACh esterase inhibitors on the end-plate potentials of frog muscle (90), together with the results of Kuffler and others, had by then convinced him that transmission at the neuromuscular junction was a chemical process mediated by ACh. He was uncertain, however, about the application of the electrical hypothesis to transmission in ganglia, and confined his restatement to monosynaptic excitation and direct inhibition in the spinal cord. He saw no need to postulate specialization of the postsynaptic membrane at excitatory synapses, and outlined the situation if Golgi cells discharged an impulse. This revised electrical hypothesis, however, could not account for the prolonged inhibition of motoneurones by impulses in cutaneous afferent fibres (92).

With colleagues including Malcolm, Brooks, CBB Downman, TH Barakan, AK McIntyre, LG Brock, W Rall, K Bradley and DM Easton, Eccles undertook a number of studies of spinal cord excitation and inhibition. These investigations set the stage for the later breakthrough intracellular experiments. He and his colleagues investigated motoneurone orthodromic and antidromic action and after-potentials (95), and synaptic potentials generated during and following repetitive excitation of low-threshold afferent fibres (103). The effects on spinal monosynaptic transmission of dorsal root section peripheral to the ganglia (115) and during chromatolysis after ventral root section (110) were also studied. The electrical thresholds, conduction velocities and central actions of impulses in Groups I, II and III muscle afferent fibres were characterized (101), and Group I fibres from thigh muscles were found to include two sub-types (Ia and Ib) (118). Both direct and poly-synaptic inhibition were found to be reduced by intravenous sub-convulsive doses of strychnine (119), an antagonism later to be of critical significance to the identification of glycine as a spinal inhibitory transmitter.

The outstanding achievement of Eccles's eight years in Dunedin, however, was undoubtedly the pioneering success he and his colleagues Brock and JS Coombs had in using microelectrodes to record intracellularly from cat spinal motoneurones in vivo. He was aware of the advances just achieved by the introduction of intracellular recording from squid giant axons and isolated muscle fibres, and wanted to use this technique in the central nervous system. Furthermore, his experimental skills and broad anatomical and physiological knowledge of the spinal cord were essential. So was his experience with recording extracellular potentials in the cat spinal cord in vivo using insulated metal electrodes. Brock developed techniques for making and filling glass microelectrodes, and Coombs, a physicist aptly described by Eccles as a 'shy genius', designed a versatile and readily operated electronic stimulating and recording unit, later widely known as the 'ESRU', together with amplifiers and a cathode-follower input stage essential for recording with high resistance electrodes.

Success came in June 1951 with the recording of resting, action and depolarizing excitatory synaptic potentials from motoneurones. The recording on 20August 1951 of membrane hyperpolarizations having time courses similar to that of direct inhibition was sensational (see 429), as the potential had the opposite polarity to that predicted by the Golgi cell hypothesis (105). Eccles immediately considered his hypothesis to have been falsified, and accepted that spinal synaptic inhibition and excitation were both chemical in nature, and mediated by two specific chemical transmitters. This rejection of electrical transmission (425, 429) was a most dramatic conversion by one of the strongest critics of chemical transmission in the mammalian central nervous system. Dale, a long-standing friend of Eccles, was later to write (1954): 'A remarkable conversion indeed. One is reminded almost inevitably of Saul on his way to Damascus when sudden light shone and the scales fell from his eyes.'

Eccles was committed to travel abroad from Dunedin in November 1951, and also expected to be without research facilities for at least a year. In 1950, concerned that the heavy teaching load in Dunedin limited his competitive edge in the rapidly advancing field of neurophysiology, he had accepted an invitation to the Chair of Physiology in the John Curtin School of Medical Research (JCSMR) in Canberra. Since no laboratories were then available in Canberra, arrangements were made for him to continue working in Dunedin after January 1951, and he took up his appointment in December that year. Meanwhile, in June 1950, the President and Fellows of Magdalen College had invited him to deliver the 1952 Waynflete Lectures. Leaving Dunedin in November 1951, Eccles flew via North America to visit colleagues and give lectures before arriving in Oxford in January 1952.

The eight Waynflete Lectures, delivered at weekly intervals, attracted large audiences. The first five dealt with basic membrane and synaptic neurophysiology, and the final three were concerned with plasticity, memory, conditioned reflexes, the cerebral cortex and the mind-brain problem. His dualistic approach to the latter, a neurophysiological hypothesis of will, first published in Nature in July 1951 (99) and elaborated further in the final Waynflete Lecture, created intense discussion. The lectures were published in 1953 as a monograph, The Neurophysiological Basis of Mind: The Principles of Neurophysiology (111), which had a considerable influence on the development of neuroscience.

In late February, Eccles visited Sherrington at a nursing home in Eastbourne. Following Sherrington's death on 4March he returned to Dunedin via the United States where he contributed to a Cold Spring Harbor Symposium on the neurone, at which there was much discussion and controversy related to intracellular recording (441). In September 1952 he and his family moved to Canberra.

Canberra

The Australian National University (ANU) and the JCSMR were established in August 1946 (Fenner 1971, Foster and Varghese 1996). Initially the Department of Physiology was located in a temporary one-storey building completed in March 1953. Eccles then began a remarkable and intense period of research activity that continued for over thirteen years. During this time, 74 investigators from 20 different countries worked in the Department (441). Of these, 41 from 14 countries collaborated and published with Eccles. He later wrote about this period: 'Without doubt it was the high point of my research career' (441), and in 1989 described it as 'my 14 golden years, scientifically speaking' (Letter to RA Hohnen, ANU Registrar during Eccles's period in Canberra). Early in 1957 the Department of Physiology moved into the permanent building of the JCSMR. The additional space, which included six large research laboratories, enabled expansion of the research staff and increased interest in neuropharmacology and neurochemistry.

With Coombs, who had accompanied him from Dunedin, and P.Fatt, Eccles began his research in Canberra with a biophysical study of the motoneurone membrane, and of synaptic excitation and inhibition in the cat lumbar spinal cord, using single and double-barrel glass intracellular electrodes (127-131). Inhibitory postsynaptic potentials (IPSPs), initially recorded as hyperpolarizations, were observed to gradually diminish and reverse to depolarizing potentials. The recognition that this was the consequence of the leakage of ions from microelectrodes containing potassium chloride led to the use of electrical currents to inject anions and cations of different hydrated ion diameter into motoneurones. Together with the first measurement of the reversal potential for IPSPs in the mammalian central nervous system, these findings suggested that an increased permeability to potassium and chloride ions occurred at the inhibitory synapses of the direct and recurrent pathways in the spinal cord (128). In contrast, excitatory postsynaptic potentials (EPSPs) were generated by a non-selective increase in the permeability to all species of ion (129), as had been demonstrated earlier for muscle end-plate potentials by Fatt and Katz (1951). Later studies by Eccles also indicated the involvement of both chloride and potassium ions in generating spinal (252) and hippocampal IPSPs (445, see 465).

Eccles, with Coombs and D.R.Curtis, analysed the antidromic, orthodromic and directly evoked action potentials of motoneurones in terms of the morphology of these cells (149, 150). Direct measurement of the specific membrane resistance and capacitance (164) enabled the time course of synaptic currents underlying EPSPs and IPSPs to be calculated (165). The rapid decline of these currents was ascribed to the diffusion of transmitter from the synaptic cleft, consistent with a theoretical analysis that Eccles published in 1957 with J.C.Jaeger (154).

Early in 1953 Eccles and Fatt (see 425) made two very significant discoveries. The first, with K.Koketsu, showed that the spinal recurrent inhibitory pathway was disynaptic. Impulses in motor axon collaterals excited, at nicotinic cholinergic synapses, interneurones which were appropriately named 'Renshaw' cells in memory of B.Renshaw who first recorded their high-frequency discharge in response to ventral root stimulation (Renshaw 1946). In turn Renshaw cells monosynaptically hyperpolarized motoneurones (123). This, the first direct evidence that ACh was a central transmitter (see 134), exemplified what Eccles called Dale's 'principle' (540), namely that the same transmitter is released at all synapses made by one neurone (Dale 1935). Renshaw cells were the first central inhibitory interneurones to be identified physiologically and pharmacologically. Later, Eccles and his colleagues recorded intracellularly from Renshaw cells, and examined their connectivity in order to elucidate the functional significance of this inhibitory mechanism (204, 205).

In 1953, direct spinal inhibition was still thought to be monosynaptic since its central latency appeared to be similar to that of the monosynaptic excitation of motoneurones. Although Brock, Coombs and Eccles had earlier found that the central latency of direct IPSPs exceeded that of monosynaptic EPSPs by as much as 1ms, this difference was ascribed to a longer intraspinal pathway of the 'inhibitory' fibres. Eccles and Fatt, with S Landgren, however, showed that direct inhibition was disynaptic (133), the inhibitory interneurones being located in the spinal intermediate nucleus. The disynaptic nature of direct inhibition was later confirmed in 1960 by Eccles, with T Araki and M Ito. Direct and recurrent inhibitions had been shown to be blocked by strychnine and Eccles, with VB Brooks and Curtis, found that tetanus toxin had a similar effect (138).

These 1953-55 investigations led Eccles to postulate that a central neurone had either an excitatory or an inhibitory action on other neurones (120, 133, 441). This proposition included inhibition of inhibitory neurones as the basis for disinhibition (see 343). Synapses of all primary afferent dorsal root fibres were excitatory, any subsequent inhibitory action was mediated by excitation of interposed inhibitory interneurones. In later investigations Eccles and his colleagues identified and established the role of other inhibitory interneurones in the spinal cord, dorsal column nuclei, the thalamus, hippocampus and cerebellum (see 343).

Eccles was invited to present the Twenty-Ninth Course of Herter Lectures at the Johns Hopkins School of Medicine, Baltimore, in October 1955. The four lectures, largely based on research carried out in Canberra, were revised for publication in 1957 as a monograph, The Physiology of Nerve Cells (137), one of the most influential books in neurobiology.

With his daughter R.M.Eccles and A.Lundberg, Eccles initiated in 1956 a series of papers on the neuronal organization within the lumbar spinal cord using intracellular recording of postsynaptic potentials, a technique that was more discriminative than the recording of neuronal discharges (208). The monosynaptic connections between Groups Ia, Ib and II afferent fibres from muscle and different types of alpha motoneurone (142, 144, 148), intermediate nucleus interneurones (191) and gamma motoneurones (187) were examined, the latter study also involving A.Iggo. He also examined the properties of chromatolysed motoneurones with B.Libet and R.R.Young (159), and the effects on monosynaptic EPSPs of peripheral section of afferent fibres with K. and R.Miledi (163). In collaboration with O.Oscarsson, Eccles also studied the cells of origin of the ventral (196) and the dorsal (197) spinocerebellar tracts, a prelude to his later investigations of the cerebellar cortex.

In 1956-1957 Eccles, with R.M.Eccles and Lundberg, discovered that significant differences existed in both the axonal conduction velocity and the after-hyperpolarization (AHP) of motoneurones innervating slow- and fast-contracting muscles. The maximum firing frequency of a motoneurone was controlled by the duration of its AHP, and matched the contraction response of its motor unit (157). Subsequently, in 1958 with A.J.Buller and R.M.Eccles, neural influences from the spinal cord were found to affect the post-natal differentiation of slow but not fast muscles in the cat hind limb (182). The crossing and subsequent regeneration of nerves to fast and slow muscles changed their contraction properties, suggesting that specific substances secreted at motoneurone axon terminals both caused and maintained differences in the contractile properties of fast and slow muscles (183).

Because of his continuing interest in plasticity (see 158), Eccles searched with R.M.Eccles and F.Magni for changes in the monosynaptic connections of cat hind limb motoneurones after various regenerations of muscle afferent nerves in kittens (190). There was, however, relatively poor synaptic plasticity of spinal cord connections in mature cats (217).

In 1960 Eccles began a comprehensive study of the mechanism and organization of what came to be called 'presynaptic' inhibition in the spinal cord. K.Frank and M.G.F. Fuortes (1957) had reported that stimulation of a flexor muscle nerve depressed monosynaptic EPSPs of extensor muscle motoneurones without recordable changes in membrane potential or excitability. The prolonged inhibitory process could be due to either a presynaptic reduction of transmitter release, or a membrane conductance increase at distal dendritic sites (Frank 1959). Eccles was in a unique position to explore the nature and significance of this type of inhibition. He had, in 1948, with C.McC.Brooks and Malcolm, observed that presynaptic spikes and excitatory synaptic potentials recorded near spinal motoneurones were reduced by a prior inhibitory input. This reduction, considered at the time to be of little physiological significance, was attributed to depolarization of excitatory presynaptic fibres (88), later to be referred to as primary afferent depolarization, PAD. Additionally, with in 1958, Eccles had recorded dorsal root potentials (DRPs) intracellularly from intraspinal afferent fibres, and also prolonged EPSPs from dorsal horn interneurones possibly involved in the generation of DRPs (179).

Over the period 1961-1965, 29 full papers dealing with various aspects of presynaptic inhibition, 21 of which Eccles co-authored, were published in refereed journals from his department, and also 13 review articles and one book (245) in which presynaptic inhibition was featured. Eccles's collaborators in studies of this inhibition in the lumbar and cervical spinal cord and dorsal column nuclei were RM Eccles, Magni, Willis, WM Kozak, RF Schmidt, PG Kostyuk, P Andersen, TA Sears, T Oshima and T Yokota. The investigations were carried out on barbituate anaesthetised cats, often cooled to accentuate the inhibition and PAD. In addition to recording DRPs, DRRs and extra- and intra-cellular potentials from intraspinal primary afferent fibres and neurones, changes in the excitability of intraspinal afferent fibres indicating PAD were determined using the extracellular microstimulating technique developed by PD Wall (1958).

Prolonged depression of monosynaptic EPSPs (and of reflexes, 212) of extensor muscle motoneurones was produced by impulses in flexor muscle Group I afferent fibres without changes in motoneurone membrane potential, excitability or EPSP time course as would be expected from a membrane conductance increase. These results suggested that the depression of monosynaptic EPSPs and reflexes was entirely a presynaptic inhibitory phenomenon (193), and PAD became synonymous with presynaptic inhibition (203).

Presynaptic inhibition, hitherto not considered a significant factor influencing spinal reflex activity, provided a negative feed-back control of sensory information into the cord and supraspinal centres. Eccles's extensive investigations (see Schmidt 1971) dealt with the organization and mechanism of PAD in the spinal cord (202, 207, 210, 211, 212, 215, 231, 232) and dorsal column nuclei (248, 268, 269). Electrical stimulation of the sensorimotor cerebral cortex also produced PAD and reduced excitatory transmission from spinal (246) and cuneate (248) afferent fibres.

PAD was considered to be generated by an increase in the ion conductance of primary afferent fibre terminals, both this increase and the depolarisation reducing the amplitude of terminal action potentials and thus affecting transmitter release (228). Eccles had proposed in 1961 that PAD may be generated by the prolonged action of a chemical transmitter at synaptic contacts on terminal boutons of afferent fibres (195). Morphological evidence for such axo-axonic synapses upon boutons in the cat spinal cord was first reported by E.G.Gray (1962). The observation with Schmidt and Willis in 1961 that picrotoxin but not strychnine reduced both PAD and the presynaptic inhibition of spinal monosynaptic reflexes led to the proposal that 4-aminobutyric acid (GABA) was the depolarizing transmitter at these axo-axonic synapses (237).

With an early exception (3), Eccles had mainly examined synaptic mechanisms in, and the organization of, the spinal cord. From late 1961, however, he concentrated upon supraspinal regions, including the somatosensory system, the hippocampus and the cerebellum. This change in direction reflected his interests in cognitive functions, and was coupled with collaboration with a number of colleagues from abroad with similar interests, including Andersen, CMc Brooks, Schmidt, Sears, Oshima, Yokota, Y Løyning, PE Voorhoeve, R Llinás, K Sasaki, P Strata, DM Armstrong, RJ Harvey and PBC Matthews. Although the effects of stimulating the cerebral cortex on transmission in the spinal cord (246), dorsal column nuclei (248) and ventrobasal thalamus (271) were examined, Eccles never studied neurones and their interconnectivity in the neo-cortex itself.

After a study of synaptic transmission and inhibitory processes in the dorsal column nuclei (269), Eccles turned to the ventrobasal complex of the thalamus (271, 272) where two interesting observations were made: a prominent recurrent postsynaptic inhibition and large post-inhibitory 'rebound' depolarizing responses with superimposed bursts of action potentials. The pivotal role of this response, labelled by Eccles 'post-anodal exaltation' (see 222), in the rhythmic and synchronized activity of thalamic cells and of various types of cortical neurone was later confirmed by others.

Taking advantage of the laminar arrangement of hippocampal synapses, Eccles's group found in 1962 that the large and prolonged chloride-dependent IPSPs recorded from pyramidal cells were generated at the soma, suggesting that basket cells, with synaptic terminals clustered around the somata of pyramidal cells, were inhibitory interneurones (253,254). This discovery served as a guide for identifying other central inhibitory neurones and synapses (see 284). The subsequent finding that cerebellar basket cells inhibited Purkinje cells through synapses located on the soma (270) led to a hypothesis that postsynaptic inhibition is largely mediated by somatic synapses (230). Later, however, cerebellar stellate cells, which synapse upon medium-size Purkinje dendrites, were also found to be inhibitory (288). Inhibition of hippocampal pyramidal cells, cerebellar Purkinje cells and ventrobasal thalamo-cortical relay cells were all found to be insensitive to strychnine (239).

Eccles's last period of experimental neuroscience, concerned with the synaptic organization and mode of operation of the cerebellum, began in Canberra in 1963 and continued in Chicago and Buffalo until his retirement from direct involvement in laboratory experimentation in 1975. In large measure due to him and his colleagues Janos Szentágothai in Budapest and Masao Ito in Tokyo, a comprehensive view of the cellular organization of the mammalian cerebellar cortex became available in the late 1960s, summarized in the influential monograph, The Cerebellum as a Neuronal Machine (317), published in 1967. It will be convenient here to give an account of Eccles's cerebellar research carried out in Canberra, Chicago and Buffalo.

In a remarkable series of letters to Nature, and subsequent detailed publications, Eccles and his collaborators described the essential properties of all major types of cerebellar neurone. Each cell type was categorized and its synaptic effect on target cells determined. Somewhat surprisingly, only the granule cells were excitatory while all other neurones were inhibitory. While Ito and his colleagues had shown that Purkinje cells monosynaptically inhibited neurones in the intracerebellar and vestibular nuclei, Eccles found that basket, stellate and Golgi cells were also inhibitory. In a series of papers inaugurating the new journal, Experimental Brain Research, for which Eccles was a founding co-editor, he observed that all of these different inhibitory interneurones had similar functional properties and could only be distinguished by their location in the cerebellar cortex (288-290).

The large inhibitory postsynaptic potentials of Purkinje cells were attributed to the activity of basket cells, terminating on Purkinje cell somata (233, 270), similar to the situation in the hippocampus. Comparing basket and Golgi cell inhibition, the latter was more focussed (0.2mm on either side of the cerebellar folia) and faster than basket cell inhibition which could spread as far as 1mm to either side (292, 312). Basket and Golgi interneurones had roughly the same threshold to parallel fibre activation. Golgi cell inhibition, however, had the lowest threshold to mossy fibre stimulation, largely due to the effective mossy fibre/Golgi cell synapses. Powerful climbing fibre excitation of Purkinje cells was elicited by stimulation of the contralateral inferior olive (255). Eccles stressed the one-to-one connectivity between one climbing fibre and a given Purkinje cell (291). Climbing fibre responses could be evoked by peripheral nerve stimulation and also occurred spontaneously.

Eccles discovered a major organizational principle by activating a thin strip of parallel fibres (288). With an ingenious de-afferented preparation, an excited strip of Purkinje cells was flanked on either side by a band of inhibition mediated by basket and stellate cells. The discovery of this arrangement became a hallmark of cerebellar cortical activation. Transmission through the mossy fibre-granule cell glomeruli was also analysed (311). Mossy fibre stimulation efficiently activated granule cells which in turn excited Purkinje cells and the three types of inhibitory neurones mentioned above (290, 316). The activity of granule cells was strongly depressed by parallel fibre activation, most likely through parallel fibre activation of Golgi cells which in turn had an inhibitory effect on granule cells. This first comprehensive analysis disclosed the mode of operation of the main cellular elements of the cerebellar cortex (311, 317, 318).

An important part of Eccles's cerebellar studies was to dissect the extra- and intra-cellular potentials recorded in the cerebellar cortex in response to activation of known afferent pathways, including the topography of the activation pattern. Using his wide experience from spinal cord work, Eccles activated specific afferent fibres from muscle, joint and skin, charting surface and depth potentials, and determined whether the afferent signals were mediated by climbing or mossy fibre inputs (335). The afferent inputs to the cerebellum showed a notable somatotopical organization, although much more widely distributed than in the somatosensory thalamo-cortical system (390). The connectivity showed a remarkable mosaic pattern and a large variation in the response sizes from different afferent nerves. A cluster of papers discuss how natural activation, like taps to the foot, produced purely excitatory responses from many mossy fibres, very similar to those elicited by electrical stimulation of the nerves (386, 387). The Purkinje cells, surprisingly, were most readily excited by impulses in cutaneous fibres, and particularly in low threshold fibres. The responses to mechanical stimulation of the skin produced by climbing fibre inputs were analysed by depressing the mossy fibre contribution with pentothal anaesthesia (384, 385). Eccles also studied the activation of intracerebellar and associated nuclei by afferent impulses in peripheral nerves. Again, impulses in cutaneous nerves of all four limbs were particularly effective (374, 382, 384, 401, 403).

Eccles was awarded a Royal Medal in 1962, and the award in 1963 of the Nobel Prize in Physiology or Medicine, shared with AL Hodgkin and AF Huxley, recognized his fundamental contributions to the ionic mechanisms of synaptic transmission in the brain. His 1964 monograph The Physiology of Synapses (245) surveyed research carried out since 1951, in his and other laboratories, on excitatory and inhibitory synapses. In the preface he acknowledged the influence on his writings of three great scientists: Ramon y Cajal, Sherrington and Dale.

Faced with retirement as Head of the Department of Physiology in 1968 at the age of 65, Eccles became concerned that the research facilities, personnel and financial support that would then be available would severely limit continuation of his research. In the absence of assured funding to support collaborators, he did not regard as acceptable a three-year appointment as a University Fellow together with his own equipment in a new laboratory in the John Curtin School. Consequently, in 1966, he resigned from the Chair of Physiology, which he had occupied since 1951, to take up an appointment as a member of the Institute of Biomedical Research, recently established by the American Medical Association in Chicago. An important factor in Eccles's decision to leave Australia was his feeling of intellectual isolation, especially in relation to his increasing interests in philosophy and the mind-brain interaction.

Chicago and Buffalo

Eccles described his period in Chicago (1966-1968) as 'the briefest, the least successful, and the most unhappy (stage) of my research career' (441, p.15). Although he established a research laboratory and continued his study of the cerebellar cortex, his understanding that adequate financial support would continue after the age of 68 failed to materialize, and problems within his group apparently created considerable dissension. Accordingly, he accepted an invitation from the State University of New York at Buffalo to establish a research unit as a Distinguished Professor of Physiology and Biophysics. His laboratories were located in temporary buildings some distance from the main university, and he described his research facilities as the best he had ever had although on a smaller scale than in Canberra (441).

In the United States, from 1966 until he retired at the end of 1975, Eccles had 20 collaborators from 11 countries including the USA, and co-authored 43 papers reporting experimental results with the following (in alphabetical order) G.I.Allen, D.S.Faber, S.T.Kitai, H.Korn, J.T.Murphy, R.A.Nicoll, L.Provini, T.Rantucci, I.Rosén, F.J.Rubia, N.H.Sabah, P.Scheid, D.W.F.Schwarz, T.Shimono, H., N.Tsukahara and T.J.Willey, and also Strata, Schmidt and Oshima who had worked with him in Canberra. Their cerebellar research has been described above, and was an important component of the very large amount of new and detailed information, including the discovery of several governing principles and cellular mechanisms, that Eccles contributed to the understanding of the cerebellum and its associated structures.

In Buffalo Eccles revisited hippocampal inhibition, the subject of his three last experimental papers. He and his colleagues reported that barbiturates prolonged postsynaptic potentials (427). He also studied the anionic permeability underlying hippocampal IPSPs, by polarization of the membrane and anion injection. The permeant anions were identical to those reported in motoneurones (444). The observations hinted at the dichotomy of hippocampal IPSPs, an early GABA-A part and a later GABA-B part, as later demonstrated by one of Eccles's coworkers on this paper, R. A. Nicoll. The Eccles group found no evidence for an outwardly directed chloride pump and concluded that an inward chloride current is the likely source for the hippocampal IPSPs (445). This was the last experimental paper from John Eccles's hand.

Switzerland

In 1975, Eccles voluntarily retired and moved to Contra in the Swiss canton Ticino, in what he described as 'idyllic mountain surroundings' (441), to dedicate himself to work on the mind-brain problem. From here he travelled extensively, attending scientific meetings, lecturing in continental Europe, the UK, Japan and North America, and playing a prominent role in the International Physicians for the Prevention of Nuclear War organization. He had visiting appointments at the University of Basle, the Max-Planck Institute for Biophysical Chemistry in Göttingen and the Max-Planck Institute for Brain Research in Frankfurt. With his books, journals and reprint collection he was able to continue his academic life, completing scientific papers and writing numerous influential reviews and books, alone and in collaboration. A particularly important concept introduced in 1978 by Eccles and PL McGeer was the recognition of two general types of the postsynaptic action of transmitters: 'ionotropic', in which the transmitter increases postsynaptic membrane conductance by directly opening ion gates, and 'metabotropic', in which the transmitter acts indirectly through intracellular metabolic reactions (465, 475).

The mind-brain problem, however, was the topic which by far occupied most of Eccles's time in the period between 1975 and his death. He wrote extensively on the subject, sought new views and explanations, and discussed the issue at numerous conferences and meetings. His final book, How the Self Controls its Brain, was published in 1994 (567).

The mind-brain problem: Philosophical considerations

Eccles recounted how, when 18 years old, he was struck by an awesome feeling of uniqueness (450, p.357). He marvelled at his own brain and its capacity for thoughts and emotions, and started a life-long search for the explanation of human achievements. Without further details, he gave this special experience as the cause for 'spending his life in the neural sciences with some continuing involvement in philosophy'. Throughout his adult life he was a declared dualist, and searched relentlessly for mechanisms by which the mind controls the body. In fact, no fewer than 18per cent of his 568 publications dealt with this issue.

Although not explicitly stated, his family's religious belief must have been important. A second reason was his own scientific curiosity and wide reading. A third, strong influence came from his mentor Sir Charles Sherrington, in particular his book Man on his Nature (1940). A final driving force may have been his scepticism towards materialism. In a letter enclosed with complimentary copies of his last book (567), Eccles wrote about scientific materialism: 'A most important program for this book is to challenge this materialism and to reinstate the spiritual self as the controller of the brain.'

Although Eccles was 'a believer in God and the supernatural' (450, p.VIII), his approach to the mind-brain problem was neither purely religious nor philosophical, but largely neurobiological with a Cartesian influence. In this respect, MacKay (1987) remarks: 'Though they [Eccles and Popper] differ in important respects from that of Descartes, they agree with him that "the brain must be open to non-physical influences if mental activity is to be effective" '. In a letter to us, BIB Lindahl (1999) concurs: 'One could say that he was a follower of Descartes. Like Descartes, Eccles's point of departure in the mind-brain field was partly religious, partly scientific, but in practice Eccles's approach was, as I see it, primarily scientific.'

Eccles first discussed mind-brain interactions in relation to voluntary actions and used the term will for the mental force he saw as the initiator (111). Later, he used mind and, later again, the term self-conscious mind. In his book The Human Psyche (482, p.2) he defined the term self-conscious mind: 'it implies knowing that one knows'. He continues: 'One can also use the term self-awareness instead of self-consciousness, but I prefer self-consciousness because it related directly to the self-conscious mind'.

Eccles searched for answers to a set of essential questions:

• how can Man's enormous capacity for thinking, memory, and emotional feeling and expression be explained?
• how can the 'Will' have such a strong and precise effect on our skeletal muscles during voluntary movement?
• since our intentions ('Will') appear so strong, can they lead to a change of brain substrates, both structurally and functionally?
• can a mind-brain interaction be localized to certain, selected parts of the brain, or even to specific cells or synapses?
• which physiological, chemical and physical processes are associated with the mind-brain interaction?

His intention was to develop testable propositions in relation to these questions. In The Self and Its Brain (450, p.355) he summarized his views on the mind-brain interaction: 'It is a very strong dualism and raises the most severe scientific problems in relationship to the interface between the world of matter-energy, in the special instance of the liaison area of the brain, and the world of states of consciousness that is referred to as the self-conscious mind. Briefly, the hypothesis states that the self-conscious mind is an independent entity that actively engages in the reading out from a multitude of active centres in the modules of the liaison areas of the dominant cerebral hemisphere.'

Eccles maintained that conscious experience is provided by the self-conscious mind by itself, and not by the neural machinery of the brain with its excitatory and inhibitory synaptic interactions (450, p.362). He further proposed that the mind-brain liaison has traffic in both directions, from the brain to the mind in perception and from mind to brain in willed action (111, p.281). His term liaison brain included all those areas of the cerebral cortex that are potentially capable of being in direct liaison with the self-conscious mind, and he located this liaison brain in the cerebral cortex of the dominant hemisphere, but only in those areas which have linguistic and ideational performance. Further, he felt that a small part, maybe less than a tenth of the cortex, in the right state of activity would be enough to give an effective mind-brain liaison (111, p.283). To illustrate the mind-brain interaction in the liaison areas, Eccles used an analogy: 'a multiple scanning and probing device that reads out from and selects from the immense and diverse patterns of activity in the cerebral cortex and integrates these selected components, so organizing them into the unity of conscious experience' (450, p.363). The language Eccles used here is similar to that used by a neuroscientist to explain neuronal interaction in an activated cortical area. He stated, however, that the self-conscious mind is not identical to some physical part of the cerebral cortex like cells or synapses.

He proposed that 'the self-conscious mind exercises a superior interpretative and controlling role upon the neural events by virtue of a two-way interaction across the interface between World1 and World2' (450, p.355), using Popper's nomenclature: World 1, the world of physical objects, and World2, the world of subjective experiences. As to possible mechanisms, he proposed: 'An attempt is made to show how the operative features of modules of the cerebral cortex can result in properties of such subtlety that they could be recipients of the weak action that are postulated to be exerted by the self-conscious mind across the interface. These actions are evident by voluntary movements as described in chapter E3 and also by the recall of memories on demand by the cognitive processes, as described in Chapter E8.' (450, p.356)

In the second-last chapter of How the Self Controls its Brain (567), and in (556), Eccles and Friedrich Beck postulated that the self-conscious mind interacts with the brain on aggregations of cortical pyramidal cell dendrites, forming structures named dendrons, and further that the self-conscious mind acts by reciprocally linking each unit of mental experience, labelled a psychon, to its specific dendron. The action of psychons was considered to involve an enhancement of the release probability of transmitter vesicles at excitatory dendritic synapses, an interaction they regarded as consistent with the laws of quantum physics.

Eccles was strongly influenced by Popper's philosophy, stemming from their contact in New Zealand in 1945. In the chemical/electrical controversy about synaptic transmission, Eccles took Popper's advice, wrote several reviews summarizing the evidence and concluded that the process was electrical (75,92). Ironically, he subsequently, in 1951, came to the opposite conclusion based upon his own intracellular recordings from spinal motoneurones. Conversely, Eccles also deeply influenced Popper. In a Festschrift article to Eccles, Popper described their first encounter as creating immediate and reciprocal sympathy, and how their common interest in the mind-brain problem made them write a book together (450), about which Popper testified: 'It became an important event in both our lives' (Creutzfeldt et al. 1984). In fact, it was Eccles who made Popper change his initial formulation to the terms World1, 2 and 3 (450, p.38). Their co-authored book The Self and its Brain (450) is by far the most cited of all Eccles's philosophical contributions.

Eccles's views on the mind-brain relationship have not been accepted by a large section of the neuroscientific community. Many opponents regarded his formulation of hypotheses as too imprecise or as untestable, and some colleagues interpreted some of the underlying experiemental observations differently from Eccles. His claim that perception is an effect of the conscious mind leaves most neuroscientists with the impression that 'the conscious mind' describes a neural entity: many do not accept that there is a distinction between the conscious mind and the activity of neurones on which it plays. There are, however, arguments in favour of Eccles's belief that mental states can influence the activity of neurones. Whatever the judgement of posterity between these two positions, Eccles deserves much credit for bringing into the open the relation between mind and brain, and for putting forward hypotheses about it which he hoped and believed would be testable.

In spite of the criticisms of his mind-brain views, there can be no doubt that he used his vast knowledge and imagination to foster real understanding. In grappling with the mind-brain problems he showed the same broad knowledge displayed in his experimental activities. He covered aspects as wide apart as conscious perception, voluntary movement, language centres, effects of brain lesions and memory functions. From this wide perspective he extracted principles of importance for his philosophical ideas. In these efforts he combined his vast knowledge with skills as a writer and lecturer.

Personality

As an individual, John Eccles combined a remarkable talent with the strong motivation and stamina necessary for an outstanding scientific career. His energy was nearly overwhelming, as was his appetite for new knowledge, particularly of the brain and its mode of operation. He was actively involved in laboratory-based research from 1927 until 1975, and was closely associated with numerous scientific collaborators and with neuroscientists in other laboratories world-wide. Eccles was indeed fortunate in being able to develop his experimental expertise during the last years of the Sherrington 'School' in Oxford, using electrophysiological stimulating and recording equipment that would now be regarded as relatively crude, and to have the opportunity to hone his skills during the next forty years of increasing technical sophistication resulting from the introduction of thermionic 'valves', cathode ray tubes, glass microelectrodes, transistors, integrated circuits and computers. In Sydney he attracted investigators of the stature of Katz and Kuffler, and later he established contact with Popper in New Zealand. Furthermore, after the pioneering achievement of recording intracellularly from motoneurones in vivo, the unrivalled opportunities and facilities provided in Canberra, including academic positions for many distinguished investigators, enabled him to exploit his new-found support of central chemical neurotransmission at both a synaptic and an organizational level, the latter interest continuing in the United States.

The following remarks apply particularly to Eccles's time in Canberra, with which we are both most familiar (D.R.C., 1954-1966; P.A., 1961-1963), and to our later encounters at meetings abroad. Colleagues from the Buffalo period confirm that the same congenial atmosphere also characterized Eccles's laboratory there. Much of his success depended on an exceptional ability to create productive research teams that were usually a blend of experienced investigators and new recruits. Often he undertook projects of interest to new arrivals in the laboratory, while some of his visitors worked on their own projects. Eccles was a prolific writer, his bibliography listing 568 items, including nineteen books of which he was the sole author of twelve. His name on a publication invariably indicated his personal participation in all aspects of the investigation. His infectious enthusiasm over a new or unexpected finding, his extensive knowledge of virtually all experimental neurophysiology, which he gladly shared, and, above all, his ideas for further experimentation were both instructive and formative for his younger colleagues. With the passage of time, the term 'Prof', used by his younger colleagues to combine respect with admiration and friendship, was replaced by 'Jack'.

As a team leader Eccles was demanding, every member of the team being expected to contribute fully in a co-operative fashion. His ambition was that his groups should make solid and substantial progress. He did, however, welcome opposition provided that the evidence for an alternative view was sound or at least reasoned. Most major investigations involved at least two long experiments each week, beginning very early in the morning and often lasting for 16-20 hours, occasionally longer and extending well into the following day. He regularly participated in the animal preparation, and took considerable pride in his anatomical knowledge and surgical expertise. He also ensured that the technical equipment available for experimentation was the best available, and insisted on having first-class electronic and mechanical workshop personnel and facilities within his own department.

A particular event appreciated by his collaborators in Canberra was a late 'tea-break' in his study, usually around 11pm. This provided an opportunity for relaxed discussion, in which he provided glimpses of his scientific life including his travels to conferences abroad, his mentors, other scientists and previous colleagues. Eccles's comments were always honest, albeit at times quite terse. Those who had made significant discoveries or other progress were, however, unreservedly praised, including his strongest competitors. Interwoven in these reminiscences was advice, including the need to remember the importance of experimental design: 'Put yourself in control of the experiment, do not let the findings run away with you!' His own experiments were always carefully planned and executed, but there was always sufficient flexibility to exploit an unexpected finding.

The mixture of scientific ambition and stimulation from Eccles as leader, coupled with the satisfaction of making interesting and significant new contributions, provided his collaborators with an experience never to be forgotten. Most established life-long friendships with Eccles and each other, as did their families, and many have made their own impact on neuroscience. To commemorate Eccles's Nobel award, 48 of his colleagues contributed brief papers to a book, Studies in Physiology, presented to John C. Eccles (Curtis and McIntyre 1965). In 1983, over 200 colleagues met in the Max-Planck Institute for Biophysical Chemistry in Göttingen to celebrate Eccles's 80th birthday, of whom 52 contributed to the commemorative volume, Sensory-Motor Integration in the Nervous System (Creutzfeldt et al. 1984). In May 1993 a similarly large number of his former colleagues and associates contributed to a Scientific Symposium held at the Max-Planck Institute of Brain Research in Frankfurt to honour Eccles's 90th birthday. Eccles gained considerable personal satisfaction from the award of a knighthood in 1958 when he was President of the Australian Academy of Science, and later, in 1990, from being appointed a Companion in the Order of Australia.

Family

John Eccles married Irene Francis Miller, whom he had met in Melbourne, on 3July 1928 in Oxford. They had five daughters and four sons, of whom their eldest daughter, Rosamond Margaret, became a neurophysiologist. She spent three years in Cambridge from 1951 as an ANU PhD scholar and subsequently was a very productive member of her father's department in Canberra from 1955 until her resignation in 1966.

In Sydney the family lived in Mosman, a short ferry trip across the harbour from the Circular Quay terminal which was close to the Sydney Hospital. Following the move to Dunedin, they also lived relatively close to the university. The acquisition in Canberra of almost a hectare of land enabled Eccles to establish an orchard and vegetable garden that were the envy of his colleagues, apart from the area of lawn which required frequent mowing. In Canberra, as earlier in Sydney and in Dunedin, he and his wife were gracious hosts, particularly at weekend tennis parties on the family court and at regular country dancing sessions for members of his staff, research students, visitors and their families. Eccles was an enthusiastic participant in both forms of exercise.

When Eccles decided to move to Chicago in 1966, his wife preferred to remain in Australia close to her family, and their marriage was dissolved in 1968. In April of that year he married Helena , a neurophysiologist who collaborated closely with him from 1966 until he ceased experimentation in 1975, after which they moved to Switzerland. Here, in a majestic mountainous landscape, Eccles concentrated on the mind-brain problem. In spite of his considerable age, he continued writing on this and related themes (for example 452, 456, 464, 467, 482, 492, 509, 515, 528, 529, 531, 535, 548, 549, 555, 556, 567). A long series of books, articles, reviews, comments, book reviews, and obituaries flowed from his hands, all hand-written, until well beyond his 90th year. In all of this activity, he was efficiently and caringly assisted by his wife Helena. Eccles's activities, however, were severely curtailed by ill health from 1994, and he died on 2May 1997 in the Hospital La Carita in Locarno. He was buried on 3May 1997, following his own wish, in Contra.

Australian Academy of Science

From 1951, Eccles was one of 23 distinguished Australian scientists, including 14 Fellows of the Royal Society of London, whose successful petitioning of the Queen led to the granting of the Charter of the Australian Academy of Science on 16February 1954 (441, see Fenner 1995). He was a member of the Provisional Council in 1953, and of the first Council of the Academy from 1954 until 1957 under the Presidency of the physicist M.L.Oliphant, and was himself President from 1957 until 1961. During his Presidency the Academy's distinctive 'dome' building (Becker House, now the Shine Dome) was constructed and opened in May 1959. Major activities of the Academy in which he was involved as President included the beginnings of the Anglo-Australian Telescope Project, the establishment of a Fauna and Flora Committee to advise on major research projects in the biological sciences, a report proposing preservation of the Kosciuszko summit area as a primitive reserve, recommendations to the Prime Minister concerning science and technology, and recommendations to the Government of a policy on oceanography. In 1963 he delivered the Academy's Matthew Flinders Lecture, established to be given as a mark of distinction by scientists of the highest standing (235).

Education

University of Melbourne, MB, BS (1925); Oxford University, BA (1927), MA, DPhil (1929).

Positions held

• 1925-1928 Rhodes Scholar, Oxford.
• 1927-1929 Christopher Welch Scholar, Oxford.
• 1927-1932 Junior Research Fellow, Exeter College, Oxford.
• 1932-1934 Staines Medical Fellow, Exeter College, Oxford.
• 1934-1937 Tutorial Fellow, Magdalen College, Oxford; University Lecturer in Physiology, Oxford.
• 1937-1943 Director, Kanematsu Memorial Institute of Pathology, Sydney Hospital, Sydney.
• 1944-1951 Professor of Physiology, University of Otago Medical School, Dunedin, New Zealand.
• 1951-1966 Professor of Physiology, John Curtin School of Medical Research, Australian National University, Canberra.
• 1966-1968 Member, Institute for Biomedical Research, American Medical Association, Chicago, Illinois, USA.
• 1968-1975 Distinguished Professor of Physiology and Biophysics, State University of New York at Buffalo, Buffalo, New York, USA.
• 1975-1997 Distinguished Professor Emeritus, State University of New York at Buffalo, New York, USA.

Honours and awards

Decorations

• 1958 Knight Bachelor. 1987 Order of the Rising Sun, Gold and Silver Stars (Japan). 1990 Companion in the Order of Australia.

Membership of learned academies and professional bodies

• 1928 Member, The Physiological Society, U.K.
• 1938 Foundation Fellow, Royal Australasian College of Physicians.
• 1941 Fellow, The Royal Society, London.
• 1950 Fellow, Royal Society of New Zealand.
• 1952 Honorary Member, The American Physiological Society.
• 1954 Foundation Fellow, Australian Academy of Science; President, 1957-1961.
• 1957 Honorary Member, Neurosurgical Society of Australasia.
• 1958 Honorary Member, Australian Association of Neurologists.
• 1959 Foreign Honorary Member, American Academy of Arts and Sciences.
• 1960 Foundation Member, Australian Physiological Society.
• 1961 Member, Pontifical Academy of Science.
• Member, Deutsche Akademie der Naturforscher Leopoldina.
• 1963 Foreign Honorary Member, Accademia Nazionale dei Lincei.
• 1964 Honorary Member, American Philosophical Society.
• Honorary Member, American Neurological Society.
• Honorary Member, Australian Physiological Society.
• Honorary Member, Société Française de Neurologie.
• 1965 Member, World Academy of Arts and Sciences.
• Honorary Member, New York Academy of Sciences.
• 1966 Foreign Associate, U.S. National Academy of Sciences.
• 1967 Honorary Member, Academia Medica Lombarda.
• Honorary Fellow, American College of Physicians.
• 1968 Honorary Fellow, Indian Academy of Sciences.
• Honorary Member, Czechoslovak Medical Society J.E. Purkyně
• 1969 Associate Member, Académie Royale de Belgique.
• 1971 Honorary Member, The Physiological Society, U.K.
• 1976 Honorary Member, European Brain and Behaviour Society.
• 1977 Honorary Member, European Neuroscience Association.
• 1982 Honorary Member, Society for Neuroscience.
• 1983 Honorary Member, Japanese Physiological Society.
• 1984 Honorary Member, Indian Physiological Society.
• Honorary Member, Australian Neuroscience Society.
• 1985 Member, Bavarian Academy of Sciences.
• 1986 Member, Academia Europoea.

Honorary degrees

• Doctor of Science: Cambridge, Tasmania, British Columbia, Marquette, Loyola, Oxford, Fribourg and Yeshiva Universities, Gustavus Adolphus College.
• Doctor of Medicine: Charles, Torino, Madrid, Ulm, Basel, Georgetown and Tsukuba Universities.
• Doctor of Laws: Melbourne University

Awards

• 1925 Rhodes Scholarship, Victoria.
• 1927 Gotch Memorial Prize, Oxford.
• 1932 Rolleston Prize, Oxford.
• 1961 Baly Medal, Royal College of Physicians.
• 1962 Royal Medal, Royal Society of London. Cook Medal, Royal Society of New South Wales.
• 1963 Cothenius Medal, Deutsche Akademie der Naturforscher Leopoldina.
• Nobel Prize in Physiology or Medicine.
• Australian of the Year.
• 1991 Cortina-Ulisse Literary Prize.
• 1993 Gold Medal of the Charles University, Prague, Czech Republic (first since 1348).

International lectureships

• 1952 Waynflete Lecturer, Magdalen College.
• 1955 Herter Lecturer, Johns Hopkins University.
• 1959 Ferrier Lecturer, The Royal Society. (delivered June 1960).
• Squibb Centenary Lecturer.
• 1963 Flinders Lecturer, Australian Academy of Science.
• Rennie Lecturer, Royal Australasian College of Physicians.
• 1965 Eddington Memorial Lecturer, University of Cambridge.
• Boyer Lectures, Australian Broadcasting Corporation.
• William G. Lennox Memorial Lecturer.
• 1966 Sherrington Lecturer, University of Liverpool.
• 1968 Alexander Forbes Lecturer, Grass Foundation.
• 1968 Dunning Trust Lecturer, Queen's University, Kingston,Ontario.
• 1969 Foerster Lecturer, University of California at Berkeley.
• 1972 Patten Memorial Lecturer, Indiana University.
• 1973 Compton Lecturer, Washington University, St.Louis.
• 1973 Phi Beta Kappa Lecturer, U.S.A.
• 1976 Pahlavi Lecturer, Iran.
• 1977 Phi Beta Kappa Lecturer, U.S.A.
• Botazzi Lecturer, Società Italiana di Fisiologia.
• 1978 Gifford Lecturer, University of Edinburgh.
• 1979 Gifford Lecturer, University of Edinburgh.
• 1980 Lecturer of 'Werner Heisenberg Vorlesungen', Carl Friedrich von Siemens Stiftung, Munich.
• 1981 Carroll Lecturer, Georgetown University.
• Lecturer, '100-Jahr-Feier Walter Rudolph Hess', Zurich.
• 1990 Idrios Lecture, Oxford.

Other marks of recognition

• 1960 Member, Research Advisory Committee, CSIRO.
• 1961 Honorary Fellow, Exeter College, Oxford.
• 1961 Kempner Visiting Professor, University of Texas Medical School.
• 1964 Honorary Fellow, Magdalen College, Oxford.
• 1966 Visiting Professor, University of California, Davis.
• 1967 Distinguished Visiting Professor, University of British Columbia.
• 1978 Visiting Professor, Department of Biology, New York University.
• 1979 Green Visiting Professor, University of Texas Medical Branch at Galveston.
• 1991 Eccles Fellowships established by Australian NHMRC.
• 1992 Eccles Lectureship established in Canberra.
• 1995 Eccles PhD Scholarships established in JCSMR.
• 1997 ANU Medical Library in the JCSMR named the Eccles Medical Sciences Library.
• 1998 The John Eccles Neuroscience Laboratory opened in JCSMR.
• 1999 Postgraduate courses at Ettore Majorana Foundation and the Centre for Scientific Culture, Erice, Sicily, Italy, named Sir John Eccles School of Neurophysiology and Neurology.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.13, no.4, 2001. A shorter version will appear in Biographical Memoirs of Fellows of the Royal Society of London, 2001. It was written by:

• David R. Curtis AC, FRS, FAA, Emeritus Professor of the Australian National University; and
• Per Andersen, Professor of Neurophysiology, Institute for Basic Medical Sciences, University of Oslo, Norway.

Acknowledgements

We are indebted to John Eccles's daughters, Dr Rosamond Mason and Mrs Mary Mennis, and to Mrs G.Mathur (Librarian, Eccles Medical Sciences Library, JCSMR), Professors F.Beck, F. Jackson, B. Libet, B.I.B. Lindahl, R.Nicoll, S.J.Redman, R.F.Schmidt and P.Strata for their assistance in the preparation of the memoir, and thank Lady (Helena) Eccles for her comments on a draft text.

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1935

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1936

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1937

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1938

52. Anticholinesterases and the rate of decay of the synaptic transmitter in the superior cervical ganglion. J.Physiol. 94, 6-7P.
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1939

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58. (with W.J. O'Connor) Action of eserine on striated muscle. J.Physiol. 95, 36-38P.
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1941

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64. (with B. Katz & S.W. Kuffler) Nature of the 'endplate potential' in curarized muscle. J.Neurophysiol. 4, 362-387.
65. (with S.W. Kuffler) Initiation of muscle impulses at neuromuscular junction. J.Neurophysiol. 4, 402-417.
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1942

68. (with B.Katz & S.W.Kuffler) Effect of eserine on neuro-muscular transmission. J.Neurophysiol. 5, 211-230.

1943

69. Synaptic potentials and transmission in sympathetic ganglion. J.Physiol. 101, 465-483.
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1944

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1945

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1946

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1947

79. Man and freedom. Twentieth Century 2, No 2, 5-23.
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1948

84. (with C.McC. Brooks & J.L. Malcolm) Responses of inhibited motoneurones. Fedn.Proc. 7, 15.
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87. (with C.McC. Brooks) Inhibitory action on a motor nucleus and focal potentials generated therein. J.Neurophysiol. 11, 401-416.
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1949

90. (with W.V. Macfarlane) Action of anti-cholinesterases on endplate potential of frog muscle. J.Neurophysiol. 12, 59-80.
91. (with T.H. Barakan & C.B.B. Downman) Electric potentials generated by antidromic volleys in quadriceps and hamstring motoneurones. J.Neurophysiol. 12, 393-424.
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1950

93. (with C.McC. Brooks & C.B.B. Downman) After-potentials and excitablity of spinal motoneurones following antidromic activation. J.Neurophysiol. 13, 9-38.
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1951

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101. (with L.G. Brock & W. Rall) Experimental investigations on the afferent fibres of muscle nerves. Proc.R.Soc.Lond.B 138, 453-475.
102. (with W.Rall) Repetitive monosynaptic activation of motoneurones. Proc.R.Soc.Lond.B 138, 475-498.
103. (with W Rall) Effects induced in a monosynaptic reflex path by its activation. J.Neurophysiol. 14, 353-376.
104. Interpretation of action potentials evoked in the cerebral cortex. Electroenceph.Neurophysiol. 3, 449-464.

1952

105. (with L.G. Brock & J.S. Coombs) The recording of potentials from motoneurones with an intracellular electrode. J.Physiol. 117, 431-460.
106. (with L.G. Brock & J.S. Coombs) Synaptic excitation and inhibition. J.Physiol. 117, 8P.
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1953

110. (with C.B.B. Downman & A.K. McIntyre) Functional changes in chromatolysed motoneurones. J.Comp.Neurol. 98, 9-36.
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114. (with J.S. Coombs & P. Fatt) The action of the inhibitory transmitter. Aust.J.Sci. 16, 1-5.
115. (with A.K. McIntyre) The effects of disuse and activity on mammalian spinal reflexes. J.Physiol. 121, 492-516.
116. (with P. Fatt & K. Koketsu) Cholinergic and inhibitory synapses in a central nervous pathway. Aust.J.Sci. 16, 50-54.
117. (with L.G. Brock & J.S. Coombs) Intracellular recording from antidromically activated motoneurones. J.Physiol. 122, 429-461.
118. (with K. Bradley) Analysis of the fast afferent impulses from thigh muscles. J.Physiol. 122, 462-473.
119. (with K. Bradley & D.M. Easton) An investigation of primary or direct inhibition. J.Physiol. 122, 474-488.

1954

120. 1954 (with P. Fatt & S. Landgren) The 'direct' inhibitory pathway in the spinal cord. Aust.J.Sci. 16, 130-134.
121. A note on Professor Sir Henry Cohen's Manson Lecture 'The status of brain in the concept of mind'. Philosophy 29, 158-159.
122. (with P. Fatt, S. Landgren & G.J. Winsbury) Spinal cord potentials generated by volleys in the large muscle afferents. J.Physiol. 125, 590-606.
123. (with P. Fatt & K. Koketsu) Cholinergic and inhibitory synapses in a pathway from motor-axon collaterals to motoneurones. J.Physiol. 126, 524-562.

1955

124. (with V.B. Brooks & D.R. Curtis) Mode of action of tetanus toxin. Nature 175, 120-121.
125. The central action of antidromic impulses in motor nerve fibres. Pflügers Arch. 260, 385-415.
126. (with R.M. Eccles & P. Fatt) The action of drugs on central cholinergic synapses. J.Physiol. 129, 40-41P.
127. (with J.S. Coombs & P. Fatt) The electrical properties of the motoneuronal membrane. J.Physiol. 130, 291-325.
128. (with J.S. Coombs & P. Fatt) The specific ionic conductances and the ionic movements across the motoneuronal membrane that produce the inhibitory post-synaptic potential. J.Physiol. 130, 326-373.
129. (with J.S. Coombs & P. Fatt) Excitatory synaptic action in motoneurones. J.Physiol. 130, 374-395.
130. (with J.S. Coombs & P. Fatt) Inhibitory suppression of reflex discharges from motoneurones. J.Physiol. 130, 396-413.
131. (with J.S. Coombs & P. Fatt) The ionic permeability of the motoneurone membrane. J.Cell.Comp.Physiol. 46, 362-363.

1956

132. (with D.R. Curtis & R.M. Eccles) Pharmacological studies on reflexes. Am.J.Physiol. 183, 606.
133. (with P. Fatt & S. Landgren) Central pathway for direct inhibitory action of impulses in largest afferent nerve fibres to muscle. J.Neurophysiol. 19, 75-98.
134. (with R.M. Eccles & P. Fatt) Pharmacological investigations on a central synapse operated by acetylcholine. J.Physiol. 131, 154-169.
135. (with P. Fatt & S. Landgren) The inhibitory pathway to motoneurones. In: Proceedings of the First International Meeting of Neurobiologists, pp.73-82.
136. (with J.S. Coombs & D.R. Curtis) Time courses of motoneuronal responses. Nature 178, 1049.

1957

137. The physiology of nerve cells. The Herter Lectures, 1955. Baltimore: Johns Hopkins Press. (Translated into Russian.)
138. (with V.B. Brooks & D.R. Curtis) The action of tetanus toxin on the inhibition of motoneurones. J.Physiol. 135, 655-672.
139. (with R.M. Eccles & A. Lundberg) Durations of after-hyperpolarization of motoneurones suppling fast and slow muscles. Nature 179, 866-868.
140. (with D.R. Curtis & R.M. Eccles) Pharmacological studies on spinal reflexes. J.Physiol. 136, 420-434.
141. Excitatory and inhibitory synaptic action. Harvey Lectures 51, 1-24.
142. (with R.M. Eccles & A. Lundberg) Synaptic actions on motoneurones in relation to the two components of the GroupI muscle afferent volley. J.Physiol. 136, 527-546.
143. The clinical significance of research work on the chemical transmitter substances of the nervous system. The Bancroft Memorial Lecture. Med.J.Aust. 1, 745-753.
144. (with R.M. Eccles & A. Lundberg) The convergence of monosynaptic excitatory afferents onto many different species of alpha motoneurones. J.Physiol. 137, 22-50.
145. The generation of impulses by nerve cells. Aust.J.Sci. 19, 161-168.
146. Tonic and phasic motoneurones and the gamma loop. Ist.Congr.Internat.Neurol., 81-87.
147. Abnormalities in the behaviour of chromatolyzed moto- neurones. Ist.Congr.Internat.Neurol., 149-150.
148. (with R.M. Eccles & A. Lundberg) Synaptic actions on motoneurones caused by impulses in Golgi tendon organ afferents. J.Physiol. 138, 227-252.
149. (with J.S. Coombs & D.R. Curtis) The interpretation of spike potentials of motoneurones. J.Physiol. 139, 198-231.
150. (with J.S. Coombs & D.R. Curtis) The generation of impulses in motoneurones. J.Physiol. 139, 232-249.
151. (with J.S. Coombs & P. Fatt) Propriétés électriques de la membrane de surface d'un motoneurone. In: Microphysiologie comparée des éléments excitables, Colloques Internationaux de CNRS 67, 73-90.
152. (with J.S. Coombs & P. Fatt) Nature du potential post synaptique inhibiteur. In: Microphysiologie comparée des éléments excitables, Colloques Internationaux de CNRS 67, 281-297.
153. Some aspects of Sherrington's contribution to neurophysiology. Notes Rec.R.Soc. 12, 216-225.
154. (with J.C. Jaeger)The relationship between the mode of operation and the dimensions of the junctional regions at synapses and motor-end organs. Proc.R.Soc.Lond.B 148, 38-56.

1958

155. (with B. Libet & R.R. Young) Responses of single chromatolysed motoneurones. Fedn.Proc. 17, 96.
156. The behaviour of nerve cells. In: The Neural Basis of Behaviour (ed. G.E.W. Wolstenholme & C. O'Connor), pp.28-47. London: Churchill.
157. (with R.M. Eccles & A. Lundberg) The action potentials of the alpha motoneurones supplying fast and slow muscles. J.Physiol. 142, 275-291.
158. Problems of plasticity and organization at simplest levels in the mammalian central nervous system. Persp.Biol.Med. 1, 379-396.
159. (with B. Libet & R.R. Young)The behaviour of chromatolysed motoneurones studied by intracellular recording. J.Physiol. 143, 11-40.
160. The physiology of imagination. Scientific American 199, 135-146.
161. (with A.J. Buller & R.M. Eccles) Controlled differentiation of muscle. J.Physiol. 143, 23-24P.
162. (with D.R. Curtis & A. Lundberg) Intracellular recording from cells in Clarke's column. Acta Physiol.Scand. 43, 303-314.

1959

163. (with K. & R.Miledi) Delayed effects of peripheral severance of afferent nerve fibres on the efficacy of their central synapses. J.Physiol. 145, 204-220.
164. (with J.S. Coombs & D.R. Curtis) The electrical constants of the motoneuronal membrane. J.Physiol. 145, 505-528.
165. (with D.R. Curtis) The time course of excitatory and inhibitory synaptic action. J.Physiol. 145, 529-546.
166. Reflections on the relationship of medical research to the physical and biological sciences. Australasian Ann.Med. 8, 95-97.
167. (with K. ) Afferent fibre potentials in the spinal cord. J.Physiol. 146, 31-32P.
168. (with A.W. Liley) Factors controlling the liberation of acetylcholine at neuromuscular junctions. Am.J.Physical Med. 38, 96-103.
169. The development of ideas on the synapse. In: The Historical Development of Physiological Thought (ed. C.McC. Brooks & P. Cranefield), pp.39-66. New York: Haffner.
170. Neuron physiology-Introduction. In: Handbook of Physiology, Neurophysiology I (ed. J. Field, H.W. Magoun & V.E. Hall), pp.59-73. Baltimore: Waverly Press.
171. Monosynaptic excitatory patterns and connections of group Ia afferent fibres from muscle. XXI Internat.Congr.Physiol.Sci.:Symposia & Special Lectures, pp.87-93.
172. (with A.J. Buller & R.M. Eccles) The influence of motoneurones on the differentiation of fast and slow muscles. XXI Internat.Congr. Physiol.Sci.: Abstracts of communications, p.45.
173. Excitatory and inhibitory synaptic action. Ann.N.Y.Acad.Sci. 81, 247-264.
174. Problems in neuropharmacology. In: Symposia et Conferences Generales, Collegium Internationale Neuro-Psycho-Pharmacologicum, pp.57-58. Amsterdam: Elsevier.
175. Plasticity at the simplest levels of the nervous system. Squibb Centenary Lectures, pp.28. New York: G.P. Putnam's Sons.
176. (with K. ) Some post-tetanic changes in primary afferent fibres. J.Physiol. 148, 22-23P.
177. (with A.J. Buller & R.M. Eccles) Further observations on the controlled differentiation of muscle. J.Physiol. 148, 78P.
178. (with D.R. Curtis) Repetitive synaptic activation. J.Physiol. 149, 43-44P.
179. (with K. ) Potential changes recorded inside primary afferent fibres within the spinal cord. J.Physiol. 149, 250-273.
180. (with K. ) Presynaptic changes associated with post-tetanic potentiation in the spinal cord. J.Physiol. 149, 274-287.

1960

181. (with D.R. Curtis) Synaptic action during and after repetitive stimulation. J.Physiol. 150, 374-398.
182. (with A.J. Buller & R.M. Eccles) Differentiation of fast and slow muscles in the cat hind limb. J.Physiol. 150, 399-416.
183. (with A.J. Buller & R.M. Eccles) Interactions between motoneurones and muscle in respect of the characteristic speeds of their responses. J.Physiol. 150, 417-439.
184. (with R.M. Eccles & F. Magni) Development of monosynaptic paths following changed motoneurone function. J.Physiol. 152, 29-30P.
185. (with W. M. Kozak & F. Magni) Dorsal root reflexes in muscle afferent fibres. J.Physiol. 153, 48-49P.
186. The properties of dendrites. In: Structure and Function of the Cerebral Cortex (ed. D.B. Tower & J.P. Shadé), pp.192-203. Amsterdam: Elsevier.
187. (with R.M. Eccles, A.Iggo & A.Lundberg) Electro- physiological studies on gamma motoneurones. Acta Physiol.Scand. 50, 32-40.
188. (with R.M. Eccles & F. Magni) Presynaptic inhibition in the spinal cord. J.Physiol. 154, 28P.
189. (with T.Araki & M.Ito) Latency of central inhibition. J.Physiol. 154, 29P.
190. (with R.M. Eccles & F. Magni) Monosynaptic excitatory action on motoneurones regenerated to antagonistic muscles. J.Physiol. 154, 68-88.
191. (with R.M. Eccles & A. Lundberg) Types of neurones in and around the intermediate nucleus of the lumbo-sacral cord. J.Physiol. 154, 89-114.
192. (with T. Araki & M. Ito) Correlation of the inhibitory postsynaptic potential of motoneurones with the latency and time course of inhibition of monosynaptic reflexes. J.Physiol. 154, 354-377.

1961

193. The nature of central inhibition. The Ferrier Lecture. Proc.R.Soc.Lond.B 153, 445-476.
194. Membrane time constants of cat motoneurones and time courses of synaptic action. Expl.Neurol. 4, 1-22.
195. The mechanism of synaptic transmission. Ergebn.Physiol. 51, 299-430.
196. (with J.I. Hubbard & O. Oscarsson) Intracellular recording from cells in the ventral spino-cerebellar tract. J.Physiol. 158, 486-516.
197. (with O. Oscarsson & W.D. Willis) Synaptic action of Group I and II afferent fibres of muscle on the cells of the dorsal spino-cerebellar tract. J.Physiol. 158, 517-543.
198. The effects of use and disuse on synaptic function. In: Brain Mechanisms and Learning (ed. J.F. Delafresnaye), pp.335-348. Oxford: Blackwells.
199. The effect of frequency of activation on transmission across synapses. In: Bioelectrogenesis (ed. C. Chagas & A. Paes-de-Carvalho), pp.297-309. Amsterdam: Elsevier.
200. Inhibitory pathways to motoneurones. In: Nervous Inhibitions (ed. E. Florey), pp. 47-86. Oxford: Pergamon.
201. The synaptic mechanism of postsynaptic inhibition. In: Nervous Inhibitions (ed. E. Florey), pp.70-86. Oxford: Pergamon.
202. (with W. Kozak & F. Magni) Dorsal root reflexes of muscle Group I afferent fibres. J.Physiol. 159, 128-146.
203. (with R.M. Eccles & F. Magni) Central inhibitory action attributable to presynaptic depolarization produced by muscle afferent volleys. J.Physiol. 159, 147-166.
204. (with R.M. Eccles, A. Iggo & A. Lundberg) Electro-physiological investigations on Renshaw cells. J.Physiol. 159, 461-478.
205. (with R.M. Eccles, A. Iggo & M. Ito) Distribution of recurrent inhibition among motoneurones. J.Physiol. 159, 479-499.
206. (with O. Oscarsson) Specificité des connections synaptique dans le système nerveux central. Actualités neurophysiologiques 3, 1-21.

1962

207. (with F. Magni & W.D. Willis) Depolarization of central terminals of GroupI afferent fibres from muscle. J.Physiol. 160, 62-93.
208. Central connections of muscle afferent fibres. In: Muscle Receptors (ed. D. Barker), pp. 81-101. Hong Kong: Hong Kong University Press.
209. (with P.G. Kostyuk & R.F. Schmidt) On the nature and functional significance of the electrotonic potential of the dorsal root of the spinal cord. J.Physiol. Kiev, 8, 21-37.
210. (with P.G. Kostyuk & R.F. Schmidt) Central pathways responsible for depolarization of primary afferent fibres. J.Physiol. 161, 237-257.
211. (with P.G. Kostyuk & R.F.Schmidt) Presynaptic inhibition of the central actions of flexor reflex afferents. J.Physiol. 161, 258-281.
212. (with R.F. Schmidt & W.D. Willis) Presynaptic inhibition of the spinal monosynaptic reflex pathway. J.Physiol. 161, 282-297.
213. (with P. Andersen & T.A. Sears) Presynaptic inhibitory action of cerebral cortex on spinal cord. Nature 194, 740-741.
214. (with P. Andersen & R.F. Schmidt) Presynaptic inhibition in the cuneate nucleus. Nature 194, 741-742.
215. (with P.G. Kostyuk & R.F. Schmidt) The effect of electric polarization of the spinal cord on central afferent fibres and on their excitatory synaptic action. J.Physiol. 162, 138-150.
216. (with R.M. Eccles & C.N. Shealy) An investigation into the effect of degenerating primary afferent fibres on the monosynaptic innervation of the spinal cord. J.Neurophysiol. 25, 544-558.
217. (with R.M. Eccles, C.N.Shealy & W.D.Willis) Experiments utilizing monosynaptic excitatory action on motoneurones for testing hypotheses relating to specificity of neuronal connections. J.Neurophysiol. 25, 559-579.
218. Bases neurophysiologiques de l'esprit.In: La vie de l'homme (ed. H. Gregoire), Chapt.4. Geneva: Rene Kister.
219. Abnormal connections in the central nervous system. In: Abnormal Nervous Function (ed. R.G. Grenell), pp.16-38. New York: Harper & Row.
220. (with R.F.Schmidt & W.D. Willis) Primary afferent depolarization of Group Ib afferent fibres of muscle. XXII Internat.Cong.Physiol. Sci: Abstracts Communicat. 919.
221. (with R.M. Eccles & W. M. Kozak) Further investigations on the influence of motoneurones on the speed of muscle contraction. J.Physiol. 163, 324-339.
222. (with P. Andersen) Inhibitory phasing of neuronal discharge. Nature 196, 645-647.
223. Spinal neurones:synaptic connexions in relation to chemical transmitters and pharmacological responses. In: Proc.Int.Pharmacol.Meeting 8 (ed. B. Uvnäs), pp.157-182. Oxford: Pergamon Press.
224. Homeostatic mechanisms in the nervous system. In: Perspectives in Biology (ed. C.F. Cori, V.G. Foglia, L.F. Leloir & S. Ochoa), pp.361-368. Amsterdam: Elsevier.

1963

225. (with R.F Schmidt & W.D. Willis) Depolarization of central terminals of Group Ib afferent fibres of muscle. J.Neurophysiol. 26, 1-27.
226. Presynaptic and postsynaptic inhibition in the central nervous system. Dai 16 Kai Nihon Igakukai Sokai Kakujitsu Koenschu IV, 95-110.
227. (with R.F. Schmidt & W.D. Willis) The location and mode of action of the presynaptic inhibitory pathways onto Group I afferent fibres from muscle. J.Neurophysiol. 26, 506-522.
228. (with R.F. Schmidt & W.D. Willis) The mode of action of the synaptic mechanism producing presynaptic inhibition. J.Neurophysiol. 26, 523-538.
229. (with I.A. Boyd) Fast- and slow-conducting small motor fibres in nerves to mammalian skeletal muscle. J.Physiol. 165, 29P.
230. (with P. Andersen & Y. Løyning) Recurrent inhibition in the hippocampus with identification of the inhibitory cell and its synapses. Nature 198, 540-542.
231. (with R.F. Schmidt & W.D. Willis) Inhibition of discharges into the dorsal and ventral spinocerebellar tracts. J.Neurophysiol. 26, 635-645.
232. (with R.F. Schmidt & W.D. Willis) Depolarization of central terminals of cutaneous afferent fibres. J.Neurophysiol. 26, 646-661.
233. (with P. Andersen & P.E. Voorhoeve) Inhibitory synapses on somas of Purkinje cells in the cerebellum. Nature 199, 655-656.
234. (with P. Andersen & Y. Løyning) Identification of inhibitory neurones in the hippocampus. Nature 199, 699-700.
235. Modes of communication between nerve cells. Fourth Matthew Flinders Lecture, Aust.Acad.Sci. Year Book 1963, pp.87-107.
236. Mind, the ultimate expression of the living state. In: The Living State, pp.34-38. Cleveland: Western Reserve University.
237. (with R.F. Schmidt & W.D. Willis) Pharmacological studies on presynaptic inhibition. J.Physiol. 168, 500-530.
238. Researches on the central nervous system. Pontificia Academia Scientiarum, Commentarii 1, 1-16.
239. (with P. Andersen, Y. Løyning & P.E. Voorhoeve) Strychnine-resistant central inhibition. Nature 200, 843-845.
240. Physiological investigations on neuromuscular transmission. Dai 16 Kai Nihon Igakuki Sokai Kakujitsu Koenshu I, 656-668.
241. Postsynaptic and presynaptic inhibitory actions in the spinal cord. In: Brain Mechanisms, Prog.Brain Res. 1, (ed. G. Moruzzi, A. Fessard & H.H. Jasper), pp.1-18. Amsterdam: Elsevier.
242. Interrelationship between nerve and muscle cell. In: The Effect of Use and Disuse on Neuromuscular Functions (ed. E. Gutman & P. Hnik), pp.19-28. Prague: Czechoslovak Academy of Sciences.
243. Specificity of neural influence on speed of muscle contractions. In: The Effect of Use and Disuse on Neuromuscular Functions (ed. E. Gutman & P. Hnik), pp.111-128. Prague: Czechoslovak Academy of Sciences.
244. Specificity of monosynaptic innervation of motoneurones. In: The Effect of Use and Disuse on Neuromuscular Functions (ed. E. Gutman & P. Hnik), pp.229-246. Prague:Czeckoslovak Academy of Sciences.

1964

245. The physiology of synapses. Berlin, Göttingen, Heidelberg: Springer-Verlag. (Translated into Japanese, Polish & Russian)
246. (with P. Andersen & T.A. Sears) Cortically evoked depolarization of primary afferent fibres in the spinal cord. J.Neurophysiol. 27, 63-77.
247. (with P. Andersen, R.F. Schmidt & T. Yokota) Slow potential waves produced in the cuneate nucleus by cutaneous volleys and cortical stimulation. J.Neurophysiol. 27, 78-91
248. (with P. Andersen, R.F. Schmidt & T. Yokota) Depolarization of presynaptic fibres in the cuneate nucleus. J.Neurophysiol. 27, 92-106.
249. (with P. Andersen & C.McC. Brooks) Electrical responses of the ventro-basal thalamus. In: Lectures on the diencephalon, Prog.Brain Res. 5 (ed. W. Bargmann & J.P. Schadé), pp.100-113. Amsterdam: Elsevier.
250. The controls of sensory communications in the brain. G.E.Rennie Memorial Lecture, 1963. Australasian.Ann.Med. 13, 102-113.
251. (with R.M. Eccles & M. Ito) Effects of intracellular potassium and sodium injections on the inhibitory postsynaptic potential. Proc.R.Soc.Lond.B 160, 181-196.
252. (with R.M. Eccles & M. Ito) Effects produced on inhibitory postsynaptic potentials by the coupled injection of cations and anions into motoneurones. Proc.R.Soc.Lond.B 160, 197-210.
253. (with P. Andersen & Y. Løyning) Location of postsynaptic inhibitory synapses on hippocampal pyramids. J.Neurophysiol. 27, 592-607.
254. (with P. Andersen & Y. Løyning) Pathway of postsynaptic inhibition in the hippocampus. J.Neurophysiol. 27, 608-619.
255. (with R. Llinás & K. Sasaki) Excitation of cerebellar Purkinje cells by the climbing fibres. Nature 203, 245-246.
256. Der Mechanismus der postsynaptischen Hemmung. Agnew Chem. 76, 674-681.
257. Inhibitory controls on the flow of sensory information in the nervous system.In: Information processing in the nervous system (ed. R.W. Gerard & J.W. Duyff), pp.24-40. Amsterdam: Excerpta Medica.
258. Editor with J.P.Schadé. Organization of the spinal cord. Prog.Brain Res. 11. Amsterdam: Elsevier.
259. The neurophysiological basis of experience. In: The critical approach to science and philosophy: Essays in honour of Karl Popper (ed. M. Bunge), pp.266-279. Glencoe: Free Press.
260. Editor with J.P. Schadé. Physiology of spinal neurones. Prog.Brain Res. 12. Amsterdam: Elsevier.
261. The excitatory responses of motoneurones. In: Physiology of spinal neurones. Prog.Brain Res. 12 (ed. J.C. Eccles & J.P. Schadé), pp.1-13. Amsterdam: Elsevier.
262. Presynaptic inhibition in the spinal cord. In: Physiology of spinal neurones. Prog.Brain Res. 12 (ed. J.C. Eccles & J.P. Schadé), pp.65-91. Amsterdam: Elsevier.
263. Neuroanatomical basis of behaviour-the ultimate units. In: Unfinished tasks in behavioural sciences (ed. A. Abrams, H.H. Garner & T.E.O. Toman), pp.12-32. Baltimore: Williams & Wilkins.
264. The ionic basis of postsynaptic inhibition. Science, N.Y.145, 1140-1147.
265. Modes of transmission within the nerve cells and between nerve cells. In: Die nervenphysiologie in gegenwartiger sicht. Nova Acta Leopoldina 28, 33-57.
266. The identification of postsynaptic inhibitory cells and synapses. Pontifica Academia Scientiarum, Commentaria 1, 44.
267. The ionic mechanism of postsynaptic inhibition. Nobel Lecture, 1963, pp.261-283. Stockholm: Nobel Foundation.
268. (with P. Andersen, R.F. Schmidt & T. Yokota) Identification of relay cells and interneurones in the cuneate nucleus. J.Neurophysiol. 27, 1080-1095.
269. (with P. Andersen, T. Oshima & R.F. Schmidt) Mechanisms of synaptic transmission in the cuneate nucleus. J.Neurophysiol. 27, 1096-1116.
270. (with P. Andersen & P.E. Voohoeve) Postsynaptic inhibition of cerebellar Purkinje cells. J.Neurophysiol. 27, 1138-1153.
271. (with P. Andersen, C.McC. Brooks & T.A. Sears) The ventro-basal thalamus: potential fields, synaptic transmission and excitability of both presynaptic and postsynaptic components. J.Physiol. 174, 348-369.
272. (with P. Andersen & T.A. Sears) The ventrobasal complex of the thalamus:types of cells, their responses and their functional organization. J.Physiol. 174, 370–399.
273. The ionic mechanisms of postsynaptic inhibition. Nobel Lecture, 1963. Aust.J.Sci. 27, 121-131.
274. (with R. Llinás & K. Sasaki) Golgi cell inhibition in the cerebellar cortex. Nature 204, 1265-1266.

1965

275. The synapse. Scientific American, 212, 56-66.
276. Pharmacology of central inhibitory synapses. Br.Med.Bull. 21, 19-25.
277. Presynaptic inhibition in the central nervous system. Acta Physiol.Hung. 26, 163-180.
278. Functional meaning of the patterns of synaptic connections in the cerebellum. Persp.Biol. Med. 8, 289-310.
279. Conscious experience and the human brain. Trans.R.Soc.N.Zealand., Gen. 1, 175-182.
280. Inhibition in thalamic and cortical neurones and its role in phasing neuronal discharges. William G.Lennox Memorial Lecture, 1964. Epilepsia, 6, 89-115.
281. Conscious experience and memory. Recent Adv.Biol.Psychiatr. 8, 235-236.
282. The control of neuronal activity by postsynaptic inhibitory action. XXIII Internat.Cong. Physiol.Sci: Invited Lectures, pp.84-95. Amsterdam: Excerpta Medica.
283. Possible ways in which synaptic mechanisms participate in learning, memory and forgetting. In: Anatomy of memory (ed. D.P. Kimble), pp.12-87. Palo Alto:Science and Behaviour Books.
284. (with P. Andersen) Locating and identifying postsynaptic inhibitory synapses by the correlation of physiological and histological data. In: Modern trends in neuro-morphology. Symp.Biol.Hung. 5, 219-242.
285. The brain and the unity of conscious experience. Eddington Memorial Lecture, 1965. Cambridge: Cambridge University Press.
286. The brain and the person. The Boyer Lectures, 1965. Sydney: Australian Broadcasting Commission.
287. (with R. Llinás & K. Sasaki) Inhibitory systems in the cerebellar cortex. Proc.Aust.Assoc. Neurologists. 3, 7-13.

1966

288. (with R. Llinás & K. Sasaki) The inhibitory interneurones within the cerebellar cortex. Expl.Brain Res. 1, 1-16.
289. (with R. Llinás & K. Sasaki) Parallel fibre stimulation and the responses induced thereby in the Purkinje cells of the cerebellum. Expl.Brain Res. 1, 17-39.
290. (with R. Llinás & K. Sasaki) The mossy fibre-granule cell relay in the cerebellum and its inhibitory control by Golgi cells. Expl.Brain Res. 1, 82-101.
291. (with R. Llinás & K. Sasaki) The excitatory synaptic action of climbing fibres on Purkinje cells of the cerebellum. J.Physiol. 182, 268-296.
292. (with R. Llinás, K. Sasaki & P. Voorhoeve) Interaction experiments on the responses evoked in Purkinje cells cells by climbing fibres. J.Physiol. 182, 297-315.
293. (with R. Llinás & K. Sasaki) The action of antidromic impulses on cerebellar Purkinje cells. J.Physiol. 182, 316-345.
294. (with R. Llinás & K. Sasaki) Intracellularly recorded responses of the cerebellar Purkinje cells. Expl.Brain Res. 1, 161-183.
295. Brain and the development of the human person. Impact, 16, 93-112. Paris: UNESCO.
296. Functional organization of the cerebellum in relation to its role in motor control. In: Muscular afferents and motor control: Nobel Symposium I (ed.R.Granit), pp.19-36. Stockholm: Almquist & Wiksell
297. Properties and functional organization of cells in the ventrobasal complex of the thalamus. In: The thalamus (ed. D.P. Purpura & M.D. Yahr), pp.129-141. New York: Columbia University Press.
298. Cerebral synaptic mechanisms. Pontifica Academia Scientiarum, 30, 41-87.
299. Conscious experience and memory. Pontifica Academia Scientiarum, 30, 469-513.
300. Editor. Brain and conscious experience. New York: Springer-Verlag.
301. Cerebral synaptic mechanisms. In: Brain and conscious experience (ed. J.C. Eccles), pp.24-58. New York: Springer-Verlag.
302. Conscious experience and memory. In: Brain and conscious experience (ed. J.C. Eccles), pp.314-344. New York: Springer-Verlag.
303. Some observations on the strategy of neurophysiological research. In: Nerve as a tissue (ed. K. Rodahl), pp.445-455. New York: Harper & Row.
304. The fundamental importance of brain research. Naturwissenschaften, 53, 165-166.
305. (with K. Sasaki & P. Strata) The profiles of physiological events produced by a parallel fibre volley in the cerebellar cortex. Expl.Brain Res. 2, 18-34.
306. Conscious experience and memory. Recent Adv.Biol.Psycht. 8, 235-256.
307. The ionic mechanisms of excitatory and inhibitory synaptic action. Ann.N.Y.Acad.Sci. 137, 473-494.
308. What is an individual? (Panel presentation with R.M. Gerard, K.M. Colby & W.R. Dennes) In: Mental health in a changing community, pp.129-134. USA: Grune & Stratton.
309. The fundamental importance of brain research. In: Conflict resolution and world education (ed. S. Mudd), pp.253-258. The Hague: Dr. W.J. Junk.

1967

310. Functional organization of the spinal cord. Anaesthesiology, 28, 31-45.
311. (with K. Sasaki & P. Strata) Interpretation of the potential fields generated in the cerebellar cortex by a mossy fibre volley. Expl.Brain Res. 3, 58-80.
312. (with K. Sasaki & P. Strata) A comparison of the inhibitory actions of Golgi cells and of basket cells. Expl.Brain Res. 3, 81-94.
313. The challenge of the brain – interview with MacKay. Science Journal, April 1967, 79-83. London: Assoc.Press.
314. The functioning of neural machinery in the central nervous system. Naturw.Rdsch.Stuttg. 20, 139-151.
315. Long-loop reflexes from muscle afferents to the brain-stem and cerebellum. Atti Acad.Med., Lombarda, 21, 158-176.
316. Excitatory and inhibitory actions of cerebellar mossy fibres on Purkinje cells. Actual. Neurophysiol.Paris, 7, 91-108.
317. (with M. Ito & J. Szentágothai) The cerebellum as a neuronal machine. New York: Springer-Verlag.
318. Circuits in the cerebellar control of movement. Proc.Natn.Acad.Sci.USA. 58, 336-343.
319. The inhibitory control of spinal reflex action. Electroenceph.Neurophysiol., Suppl. 25, 20-34.
320. (with L. Provini, P. Strata & H. ) Patterns of climbing fibre input to the cerebellar anterior lobe. Brain Res. 5, 425-430.
321. Evolution and the conscious self. In: The human mind (ed. J.D. Roslansky) pp.3-28. Amsterdam: North Holland.
322. Postsynaptic inhibition in the central nervous system. In: The neurosciences (ed. G.C. Quarton, T. Melnechuk & F.O. Schmitt), pp.408-427. New York: Rockefeller University Press.
323. Neurosciences, achievement and promise. The Sciences, 7, 16-19.
324. Book review of 'Of molecules and men', by Frances Crick. Zygon, 2, 281-83.

1968

325. The effect of nerve cross union on muscle contraction. In: Exploratory concepts in muscular dystrophy and related disorders, pp.151-163. Amsterdam: Excerpta Medica.
326. (with D.M. Armstrong. R.J. Harvey & P.B. Matthews) Responses in the dorsal accessory olive of the cat to stimulation of hind limb afferents. J.Physiol. 194, 125-145.
327. Professor Ragnar Granit's contributions in the field of motor control. Am.J.Phys.Med. 47, 3-7.
328. Experiméntation sur l'homme en neurophysiologie. Table ronde: La science biomedical devant le dileme de l'expérimentation sur l'homme. p.19. Paris: UNESCO.
329. The way in which the cerebellum processes sensory information from muscle. In: Neurophysiological basis of normal and abnormal motor activities (ed. M.D. Yahr & D.P. Purpura), pp.379-406. New York: Raven Press.
330. Postsynaptic inhibition in the central nervous system. In: Structure and function of inhibitory neuronal mechanisms (ed. C. von Euler, S. Skoglund & U. Söderberg), pp.291-308. Oxford: Pergamon Press.
331. Integration of information in the cerebellum. Proc. Internat.Union Physiol.Sci.XXIV Cong. 6, 19-20.
332. (with H. , L. Provini & P. Strata) Fine patterns of projection from hindlimb afferents to the cat cerebellar anterior lobe. Proc.Internat. Union Physiol. Sci. XXIV Cong. 7, 326.
333. Two hitherto unrecognized publications by Sir Charles Sherrington, O.M., F.R.S. Notes Rec.R.Soc.Lond. 23, 86-100.
334. (with L. Provini, P. Strata & H. ) Analysis of electrical potentials evoked in the cerebellar anterior lobe by stimulation of hindlimb and forelimb nerves. Expl.Brain Res. 6, 171-194.
335. (with L. Provini, P. Strata & H. ) Topographical investigations on the climbing fibre inputs from forelimb and hindlimb afferents to the cerebellar anterior lobe. Expl.Brain Res. 6, 195-215.
336. The importance of brain research for the educational, cultural and scientific future of mankind. Persp.Biol.Med. 12, 61-68.
337. (with H. & N.Tsukahara) Excitation and inhibition of cells in the cerebellum of Mustelus canis. Biol.Bull. 135, 418.
338. (with N. Tsukahara & H. ) The responses of granule cells in the cerebellum of Mustelus canis. Biol.Bull. 135, 440.
339. (with S.T. Kitai, H. & N. Tsukahara) The distribution to the cerebellar anterior lobe of climbing and mossy fibre inputs from plantar and palmar cutaneous afferents. Expl.Brain Res. 7, 1-10.
340. Influence of spino-cerebellar pathways on the cerebellar control of spinal motoneurones. Electroenceph.Neurophysiol. 25, 394.

1969

341. Central cholinergic transmission and its behavioural aspects. Historical introduction. Fedn.Proc. 28, 90-94.
342. The coordination of information in the cerebral cortex. In: Modern neurology papers in tribute to Denny-Brown (ed. S.Locke), pp.135-147. Boston: Little, Brown & Co.
343. The inhibitory pathways of the central nervous system. The Sherrington Lectures, IX, 1966. Liverpool: Liverpool University Press.
344. (with D.S. Faber, J.T. Murphy. N.H. Sabah & H. ) Firing patterns of Purkinje cells in response to volleys from limb nerves. Brain Res. 14, 222-226.
345. The future of brain sciences. In: The future of the brain sciences (ed. S. Bogoch), pp.xxxiii-xxxix. New York: Plenum Press.
346. The experiencing self. In: The Human Mind (ed. J.D. Roslansky), pp. 2-28. Amsterdam: North-Holland Publishing.
347. (with H. Korn, H. & N. Tsukahara) Slow potential fields generated in cerebellar cortex by mossy fibre volleys. Brain Res. 15, 276-280.
348. The necessity of freedom for the free-flowering of science. Dunning Trust Lecture, Queens University. Queens Quarterly, Kingston, Ontario.
349. Excitatory and inhibitory mechanisms in the brain. In: Basic mechanisms of the epilepsies (ed. H. Jasper, A. Ward & A. Pope), pp.229-261. Boston: Little, Brown & Co.
350. The development of the cerebellum of vertebrates in relation to the control of movement. Naturwissenschaften, 56, 525-534.
351. The topography of the mossy and climbing fibre inputs to the anterior lobe of the cerebellum. In: The cerebellum in health and disease (ed. W.S. Fields & W. Willis, Jr.), pp.231-262. St Louis: Warren H. Green.
352. The dynamic loop hypothesis of movement control. In: Information processing in the nervous system (ed. K.N. Leibovic), pp.245-269. Berlin: Springer-Verlag.

1970

353. (with H. & N. Tsukahara) Responses of the cells of a selachian cerebellum (Mustelus canis). Brain Res. 17, 57-86.
354. (with H. & N. Tsukahara) Responses of the granule cells of the selchian cerebellum (Mustelus canis). Brain Res. 17, 87-102.
355. (with D.S. Faber, S.T. Kitai, T. Shimono & H. ) Inhibition of antidromic invasion of cells in the selachian cerebellum. Brain Res. 17, 360-365.
356. Alexander Forbes and his achievements in electrophysiology. Persp.Biol.Med. 13, 388-404.
357. Neurogenesis and morphogenesis in the cerebellar cortex. Proc.Natn.Acad.Sci.USA 66, 294-301.
358. (with D.S. Faber, J.T. Murphy, N.H. Sabah & H. ) The integrative performance of the cerebellar cell. In: Excitatory synaptic mechanisms (ed. P. Andersen & J.K.S.Jansen), pp.223-236. Oslo: Oslo University Press.
359. Facing reality: Philosophical adventures of a brain scientist. New York: Springer-Verlag. (Translated into German, Spanish, Italian).
360. Some implications for the scientiae for the future of mankind. Studium Generale, 23, 917-924.
361. (with D.S. Faber & H. ) Antidromic invasion of cerebellar cells. J.Physiol. 210, 173P.

1971

362. (with D.S. Faber & H. ) The action of a parallel fibre volley on the antidromic invasion of cells of the cerebellum. Brain Res. 25, 335-356.
363. Animal experimentation versus human experimentation. In: Defining the laboratory animal. IVth Symposium, Internat.Committee on Laboratory Animals, pp.285-293. Washington D.C.: Natl.Acad.Sci.USA.
364. (with N.H. Sabah, R.F. Schmidt & H. ) Cutaneous mechanoreceptors acting on cerebellum via climbing fibres. Fedn.Proc. 30, 664.
365. (with N.H. Sabah, R.F. Schmidt & H. ) Cerebellar cell responses to cutaneous mechanoreceptors. Brain Res. 30, 419-424.
366. (with D.S. Faber, J.T. Murphy, N.H. Sabah & H. ) Afferent volleys in limb nerves influencing impulse discharges in cerebellar cortex. I. In mossy fibres and granule cells. Expl.Brain Res. 13,15-35.
367. (with D.S. Faber, J.T. Murphy, N.H. Sabah & H. ) Afferent volleys in limb nerves influencing impulse discharges in cerebellar cortex. II. In cells. Expl.Brain Res. 13, 36-53.
368. (with D.S. Faber, J.T. Murphy, N.H. Sabah & H. ) Investigations on integration of mossy fibre inputs to cells in the anterior lobe. Expl.Brain Res. 13, 54-77.
369. Neurobiology:the new wave leading on from molecular biology. Fedn.Proc. 30, 1413-1415.
370. The cell: its physiological properties and performance. In: Jan Evangelista Purkynê, 1787-1869. Centenary Symposium, Prague, 1969 (ed. V.Kruta), pp.151-165. Brno: Bruno University.
371. Physiological significance of the cutaneous mechanoreceptor input to the cerebellum. Proc.Internat.Union Physiol.Sci., XXV Cong. p.156.
372. Physiologie der Nervenzelle und ihrer Synapsen. In: Allegmeine Neurophysiologie, Band 10, Physiologie des Menschen. pp.108-148. Munich: Urban & Schwarzenberg.
373. (with N.H. Sabah, R.F. Schmidt & H. Táboříková) Significance of dual input to cat cerebellum via mossy and climbing fibres. J.Physiol. 218, 90-91P.
374. (with N.H. Sabah & H. ) Responses evoked in neurons of the fastigial nucleus by cutaneous mechanoreceptors. Brain Res. 35, 523-527.
375. Functional significance of arrangements of neurones in cell assemblies. Arch.Psychiatr. Nervenkr. 215, 92-106.
376. Jan-Friedrich Tönnies. J.Neurophysiol. 34, 937.
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1972

378. Editor with A.G. Karczmar. Brain and human behaviour. Heidelberg: Springer-Verlag.
379. Possible synaptic mechanisms subserving learning. In: Brain and human behaviour (ed. A.G. Karczmar & J.C. Eccles), pp.39-61. Heidelberg: Springer-Verlag.
380. (with N.H. Sabah, R.F. Schmidt & H. ) Mode of operation of the cerebellum in the dynamic loop control of movement. Brain Res. 40, 73-80.
381. The role of the cerebellum in controlling movement. In: Modern trends in physiology (ed. C.B.B. Downman), pp.86-111. London: Butterworth.
382. (with I. Rosén, P. Sheid & H. ) Cutaneous afferent responses in interpositus neurones of the cat. Brain Res. 42, 207-211.
383. Unconscious actions emanating from human cerebral cortex. Philosophic Exchange, 1, 249-252.
384. (with N.H. Sabah, R.F. Schmidt & H. ) Cutaneous mechanoreceptors influencing impulse discharges in cerebellar cortex. I. In mossy fibres. Expl.Brain Res. 15, 245-260.
385. (with N.H. Sabah, R.F. Schmidt & H. ) Cutaneous mechanoreceptors influencing impulse discharges in cerebellar cortex. II. In cells by mossy fibre input. Expl.Brain Res. 15, 261-277.
386. (with N.H. Sabah, R.F. Schmidt & H. ) Cutaneous mechanoreceptors influencing impulse discharges in cerebellar cortex. III. In cells by climbing fibres. Expl.Brain Res. 15, 484-497.
387. (with N.H. Sabah, R.F. Schmidt & H. ) Integration by cells of mossy and climbing fibre inputs from cutaneous receptors. Expl.Brain Res. 15, 498-520.
388. Ideology and motivation: an attempt towards an anthropological prognosis. Universitas, 10, 1023-1029.
389. Specific ionic conductances at synapses. In: Perspectives in membrane biophysics, a tribute to Kenneth S. Cole (ed. D.P. Agin), pp.219-243. London: Gordon & Breach.

1973

390. 1973 The cerebellum as a computer: patterns in space and time. J.Physiol. 229, 1-32.
391. A re-evaluation of cerebellar function in man. In: New developments in electromyography and clinical neuro-physiology. Vol.3 (ed. J.E.Desmedt), pp.209-224. Basel: Karger.
392. Brain, speech and consciousness. Naturwissenschaften 60, 167-176.
393. The discipline of science with special reference to the neurosciences. Daedalus 102, 85-99.
394. (with P. Scheid & H. ) Responses of red nucleus neurones to cutaneous afferent inputs. Brain Res. 53, 440-444.
395. The understanding of the brain. New York: McGraw-Hill (Translated into Spanish, German, Italian, Japanese, Mexican).
396. Reflexions on mind. Review of The nature of mind by A.J.P. Kenny, H.C. Longuet-Higgins, J.R. Lucas & C.H. Waddington. Nature 244, 183-184.
397. Brainwaves. Review of Explorers of the brain by L.A.Stevens. Nature 246, 363-364.
398. Trophic influences in the mammalian central nervous system. In: Development and aging in the nervous system (ed. M.Rockstein), pp.89-103. New York: Academic Press.
399. Cultural evolution versus biological evolution. Zygon 8, 282-293.
400. How conscious is the brain? Review of The conscious brain by S.Rose. Times Higher Educ.Suppl. 102, 15.
401. (with R.A. Nicoll, D.W.F. Schwarz & H. ) Cerebellospinal pathway via the fastigial nucleus and the medial reticular formation. Brain Res. 66, 525-530.

1974

402. Our brains and our future. In: The greatest adventure (ed. E.H. Hone & H.J. Jordon), pp.70-83. New York: Rockefeller University Press.
403. (with N.H. Sabah & H. ) Excitatory and inhibitory responses of neurones of the cerebellar fastigial nucleus. Expl.Brain Res. 19, 61-77.
404. (with N.H. Sabah & H. ) The pathways responsible for excitation and inhibition of fastigial neurones. Expl.Brain Res. 19, 78-99.
405. (with T. Rantucci, N.H. Sabah & H. ) Somatotopic studies on cerebellar fastigial cells. Expl.Brain Res. 19, 100-118.
406. Trophic interactions in the mammalian central nervous system. In: Trophic functions of the neuron (ed. D.B.Drachman), Ann.N.Y.Acad. Sci. 228, 406-423.
407. The world of objective knowledge. In: The philosophy of Karl Popper (ed. P.A. Schilpp), pp.349-370. The Library of Living Philosophers, XIV. La Salle, Ill: Open Court Publishing.
408. The physiology and physics of the free will problem. In: Progress in the neurosciences and related fields (ed. S.L. Mintz & S.M. Widmayer), pp.1-40. New York: Plenum Press.
409. Culture: The creation of man and the creator of man. In: Modern science and moral values. Proc.2nd.Internat.Conf. on the Unity of the Sciences, Tokyo, pp.23-62. New York: International Cultural Foundation.
410. Falsifiability and freedom. Debate with Sir Karl Popper. In: Reflexive water (ed. F. Elders), pp.69-131. London: Souvenir Press.
411. (with I. Rosén. P. Scheid & H. ) Temporal patterns of responses of interpositus neurons to peripheral afferent stimulation. J.Neurophysiol. 37, 1424-1437.
412. (with I. Rosén, P. Scheid & H. ) Patterns of convergence onto interpositus neurons from peripheral afferents. J.Neurophysiol. 37, 1438-1448.
413. (with T. Rantucci, I. Rosén, P. Scheid & H. ) Somatotopic studies on cerebellar interpositus neurons. J.Neurophysiol. 37, 1449-1459.
414. (with P. Scheid) Physiologie der Nervenzelle und ihrer Synapsen. In: Allgemeine Neurophysiologie, 2nd edn., vol. 10, Physiologie des Menschen (ed. O.H.Gauer, K.Kramer & R.Jung), pp. 109-64. Munich: Urban & Schwarzenberg.
415. Cerebral activity and consciousness.In: Studies in the philosophy of biology (ed. F.J.Ayala & T.Dobzhansky), pp.87-107. London: Macmillan Press.

1975

416. Hierarchical concepts in the control of movement. In: The creative process in science and medicine (ed. H.A. Krebs & J.H.Shelley), pp.61-85. Amsterdam: Excerpta Medica.
417. (with R.A. Nicoll, D.W.F. Schwartz, H. & T.J. Willey) Reticulospinal neurons with and without monosynaptic inputs from cerebellar nuclei. J.Neurophysiol. 38, 513-530.
418. (with R.A. Nicoll, H. & T.J. Willey) Medial reticular neurons projecting rostrally. J.Neurophysiol. 38, 531-538.
419. Some aspects of the dialogue between biologists and philosophers in the 20th century: Communication systems of the brain. Académie Royale de Belgique: Connaissance scientifique et philosophie, pp 251-290. Brussels: L'Académie Royale de Sciences, letters et des beaux- arts de Belgique.
420. (with I. Rosén, P. Scheid & H. ) The differential effect of cooling on the responses of the cerebellar cortex. J.Physiol. 249, 119-138.
421. (with P. Scheid & H. ) Responses of red nucleus neurones to antidromic and synaptic activation. J.Neurophysiol. 38, 947-964
422. (with T. Rantucci, P. Scheid & H. ) Somatopic studies on red nucleus: spinal projection level and respective receptive fields. J.Neurophysiol. 38, 965-980.
423. (with N.H. Sabah, R.F. Schmidt & H. ) A study of the cutaneous mechanoreceptive modalities projecting to the cerebellum. In: The somatosensory system (ed. H.H. Kornhuber), pp.125-134. Stuttgart: Thieme.
424. Letters from C.S. Sherrington, F.R.S. to Angelo Ruffini between 1896 and 1903. Reprinted from Notes Rec.Roy.Soc.Lond. 30, 69-88.
425. Under the spell of the synapse. In: The neurosciences: paths of discovery (ed. F.G. Worden, J.P. Swarzey & G. Adelman), pp.159-179. Cambridge, Mass.: MIT Press.
426. (with P. Scheid) Music and speech: artistic functions of the human brain. Psychology of Music 3, 21-35.
427. (with R.A. Nicoll, T. Oshima & F. Rubia) Prolongation of hippocampal inhibitory postsynaptic potentials by barbiturates. Nature 258, 625-627.

1976

428. The plasticity of the mammalian central nervous system with special reference to new growths in response to lesions. Naturwissenschaften 63, 8-15.
429. From electrical to chemical transmission in the central nervous system. Notes Rec.Roy.Soc. Lond. 30, 219-230.
430. (with R.A. Nicoll, D.W.F. Schwarz, H. & T.J. Willey) Medial reticular and perihypoglossal neurons projecting to cerebellum. J.Neurophysiol. 39, 102-108.
431. (with R.A. Nicoll, T. Rantucci, H. & T.J. Willey) Topographic studies on medial reticular nucleus. J.Neurophysiol. 39, 109-118.
432. The coding of stimulus intensity by the frequency of impulse discharges from receptors. Trends Biochem.Sci. 1, 68.
433. Keynote address. In: The centrality of science and absolute values. pp.1-5. New York: International Cultural Foundation.
434. Hemispheric function in the human brain. Persp.Biol.Med. 19, 290-293.
435. Brain and free will. In: Consciousness and the brain (ed. G.G. Globus, G. Maxwell & I. Savodnik), pp.101-121. New York: Plenum Press.
436. How dogmatic can materialism be? In: Consciousness and the brain (ed. G.G. Globus, G. Maxwell & I. Savodnik), pp.155-60. New York: Plenum Press.
437. The brain-mind problem as a frontier of science. Prog.Sci.Culture 1, 7-17.
438. The three joint winners of the 1975 Nobel Prize in Medicine. Prog.Sci.Culture 1, 93-96.
439. Cerebellar cortex: Chairman's remarks. In: Afferent and intrinsic organization of laminated structures in the brain. Expl.Brain Res., Suppl. 1, 3-7.

1977

440. Brain-mind problem as a frontier of science. In: The future of science: 1975 Nobel Conference (ed. T.C.L. Robinson), pp.73-104. New York: Wiley.
441. My scientific odyssey. A.Rev.Physiol. 39, 1-18.
442. An instruction-selection theory of learning in the cerebellar cortex. Brain Res. 127, 327-352.
443. (with P. Buser) Paul Dell (1915-1976). Expl.Brain Res. 27, 233-235.
444. (with G.I. Allen, R.A. Nicoll, T. Oshima & F.J. Rubia) The anionic permeability of the inhibitory postsynaptic membrane of hippocampal pyramidal cells. Proc.R.Soc.Lond.B 198, 345-361.
445. (with G.I. Allen, R.A. Nicoll, T. Oshima & F.J. Rubia) The ionic mechanisms concerned in generating the i.p.s.p.s of hippocampal pyramidal cells. Proc.R.Soc.Lond.B 198, 363-384.
446. Chairman's address. In: The search for absolute values: harmony amongst the sciences. Proc.5th.ICUS Meeting, pp.13-18. New York: International Cultural Foundation.
447. Objectivity in the neurobiological sciences: the dialogue between philosophers on the mind-brain problem. In: The case of objectivity. Proceedings of the 3rd. International Symposium at Athens & Pelion, pp.295-305. Athens: Hellenic Soc. Humanistic Studies.
448. Reflections on a life time of experiences in scientific publications. In: Semper attentus. Beiträge für Heinz Götze zum 8 August 1977, pp.95-102. Berlin: Springer-Verlag.
449. Cerebellar function in the control of movement (with special reference to the pioneering work of Sir Gordon Holmes). In: Physiological aspects of clinical neurology (ed. F.C. Rose), pp.157-178. Oxford: Blackwell Scientific Publications.
450. (with K.R. Popper) The self and its brain. An argument for interactionism. Berlin, New York: Springer International. (Translated into Spanish, German, Italian, Portuguese).
451. Evolution of the brain in relation to the development of the self-conscious mind. Ann.N.Y.Acad.Sci. 299, 161-179.
452. Hirn und Bewusstsein. In: Mannheimer Forum, 77-78, pp.9-63. Mannheim: Boehringer.
453. Learning and memory in human brain and conscious mind. Hexagon, 5. No.8, 1-10. Basel: Hoffmann-La Roche.
454. Edgar Douglas Adrian, 1889-1977. Expl.Brain Res. 31, 153-154.

1978

455. An instruction-selection hypothesis of cerebral learning. In: Cerebral correlates of conscious experience (ed. P.A. Buser & A. Rougel-Buser), pp.155-175. Amsterdam: Elsevier.
456. A critical appraisal of brain-mind theories. In: Cerebral correlates of conscious experience (ed. P.A. Buser & A. Rougel-Buser), pp.347-355. Amsterdam: Elsevier.
457. An approach to neurobiology. Review of The purposive brain by Ragnar Granit. Persp.Biol. Med. 21, 466-469.
458. Another approach to neurobiology. Review of Local circuit neurones by Pasko Rakic. Persp. Biol.Med. 21, 469-470.
459. Climbing fibre inputs from the interosseus and knee joint nerves. In: Studies in neurophysiology, presented to A.K. McIntyre (ed. R. Porter), pp.105-108. Cambridge: Cambridge University Press.
460. Lo sviluppo della persona umana: il ruola della deprivazione e della sofferenza. In: L'Uomo della sindone. Roma: Orizzonte Medica.
461. Chairman's addresses. In: The search for absolute values in a changing world. Proc.6th. ICUS Meeting, pp.13–16 & 1271-1272. New York: International Cultural Foundation.
462. The evolution of altruism. In: The search for absolute values in a changing world. Proc.6th.ICUS Meeting, pp.827-831. New York: International Cultural Foundation.
463. A critical appraisal of brain-mind problems. In: The search for absolute values in a changing world.Proc.6th. ICUS Meeting, pp.961-966. New York: International Cultural Foundation.
464. Das Problem von Gehirn und Geist. In: Die psychologie des 20 jahrhunderts, pp.1131-1177. Zurich: Kindler-Verlag.
465. (with P.L. McGeer & E.G. McGeer) Molecular neurobiology of the mammalian brain. New York, London: Plenum Press.
466. (with W. Feindel) Wilder Graves Penfield (1891-1976). Biogr.Mems.Fellows Roy.Soc.Lond. 24, 472-513.

1979

467. The human mystery. The Gifford Lectures, 1977-78. Berlin: Springer-Verlag. (Translated into Italian, German, French)
468. (with W.C. Gibson) Sherrington, his life and thought. Berlin, New York: Springer International.
469. Creation of the self. In: The early development of the affects and moods. Bull.Menninger Clinic, 43, 3-19.
470. Synaptic plasticity. Naturwissenschaften 66, 147-153.
471. Der Nervenapparat des Gehirn und des Wissen und Macht. In: Europaisches Forum, Alpbach 1978 (ed. O.Molden), pp.90-107. Wien: F.Molden Verlag.
472. Two approaches to neurobiology. Bioscience 28, 204.
473. Introductory remarks. In: Cerebro-cerebellar interactions (ed. J.Massion & K.Sasaki), pp.1-18. Amsterdam: Elsevier.
474. Hippocampal plasticity. In: Development and clinical specificity of neurons. Prog.Brain Res. 51, (ed. M. Cuenod, G.W. Kreutzberg & F.E. Bloom), pp.133-138. Amsterdam: Elsevier.
475. (with P.L. McGeer) Ionotropic and metabotropic synaptic transmission. Trends Neurosci. 2, 39-40.
476. The human brain and the human person. In: The re-evaluation of existing values and the search for truth. Proc.7th.ICUS Meeting, pp.1125-1140. New York: International Cultural Foundation.
477. Nei meandri del cervello alla scoperta delle mente. Scienza e Vita nuova, 1, 18-25.
478. The human person and the brain. Civilta delle Macchine Diretta da Francesca d'Arcais, Anno 26, Nr 1-6, 20-26.
479. La realizzazione del 'Se'. Revista Medica della Suizzera Italiano, Dec.1979, 537-544.

1980

480. The discipline of biological science with special reference to the neurosciences. Riv.Biologica, 73, 11-20.
481. Altruisms und pseudaltruisms. In: Der Mensch in der unvollkommen Gesellschaft. Europäisches Forum Alpbach 1979 (ed. O. Molden), pp.72-81. Wien: F.Molden.
482. The human psyche. The Gifford Lectures, 1979. Berlin: Springer-Verlag. (Translated into Italian, German)
483. (with H. Zeier) Gehirn und Geist. München: Kindler.
484. Event-related potentials in the cerebral cortex. In: Nerve cells, transmitters and behaviour (ed. R. Levi-Montalcini), pp.551-561. Vatican City: Pontifica Academia Scientiarum.
485. Brain, freedom and moral responsibility. Proceedings 8th. Internat Conf. Unity Sciences, pp.147-155. New York: International Cultural Foundation.
486. The emotional brain. Bull.Mem.Acad.R. Med.Belg.125, 697-713.
487. Review of Evolution and consciousness: Human systems in transition, ed. by E. Jantsch & C.H. Waddington. Zygon 15, 438-440.
488. Values and decisions. Review of The biological origins of human values by G.E. Pugh. Higher Educ.Rev. 12, 84-87.
489. New text asserts psychobiology as an independent science. Review of The psychobiology of mind by W.R. Uttal. Bioscience, 29, 482–483.

1981

490. Modular organization principles in the central nervous system. In: Regulatory functions of the CNS subsystems. Advances in Physiological Sciences, 2, 55-57. Budapest: Akademia Kiado.
491. In praise of books. Für Klaus Pipper zum 70 Geburtstag, pp.48-50. München: Pipper & Co. Verlag.
492. The self-conscious mind and the meaning and mystery of personal existence. Teachers College Record, 82, No.3, 403-426.
493. Die Menschliche Personlichkeit: ein wissenschaftliches und philosophisches Problem. Naturw.Rdsch.Stuttg. 34, 227-237.
494. The modular operation of the cerebral neocortex considered as the material basis of mental events. Neuroscience, 6, 1839-1856.
495. Physiology of motor control in man. Appl.Neurophysiol. 44, 5-15.
496. Review of Programs of the brain by J.Z. Young. Quart.Rev.Biol. 54, 361.
497. More on the body-mind problem. Review of The brain's mind: A neuroscience perspective on the mind-body problem, ed. by D. Bendra. Trends Neurosci. 4, No.9, xx.
498. The story of the International Neurobiology Society. IBRO News 9. No.2, 1-3.
499. Obituary: Derek Denny-Brown (1901-1981). IBRO News 9, No.4, 9-10.
500. Sprache, Denken und Gehirn. In: Kindler's Enzyklopädie 'Der Mensch', vol.6, pp.275-304. München: Kindler.
501. Mente. In: Enciclopadie del novecento, 4, 91-102. Instituto Enciclopedie Italiana.
502. Voluntary movement. Freiburger Universitätsblätter, 74, 24-28.
503. Language, thought and brain. Epistemologia 4, 97-126.

1982

504. Cerebral mechanisms in pain perception. Panminerva Med. 24, 1-13.
505. Life in Sherrington's laboratory, his last decade in Oxford (1925-1935). Trends Neurosci 5, 108-110.
506. The synapse: from electrical to chemical transmission. A.Rev.Neurosci. 5, 325-339.
507. The initiation of voluntary movements by the supplementary motor area. Arch.Psychiatr. Nervenkr. 231, 423-441.
508. Review of Organization in the spinal cord by A.G.Brown. Naturwissenschaften, 69, 560.
509. The liaison brain for voluntary movement:the supplementary motor area. Acta Biol.Acad. Sci.Hung. 33, 157-172.
510. Some circuit operations in the mammalian brain. Acta Anat. 113, 325-339.
511. The future of studies on the cerebellum. Expl.Brain Res.Suppl. 6, 607-620.
512. Review of Chemical transmission – 75 years, by L. Stjärne, P. Hedqvist, H. Lagercrantz & A. Wennmalm. Neuroscience 7, 2561-2562.
513. Bewusstsein der Tiere und Ich-bewusstsein des Menschen: Ein Ratsel der Evolution. Naturw.Rdsch.Stuttg. 35, 393-399.
514. Our fragile brains. Review of A Christian perspective in brain research by D.G. Jones. Zygon 17, 219-223.
515. My living dialogue with Popper. In: In pursuit of truth. Essays in honour of Karl Popper's 80th birthday (ed. P. Levinson), pp.221-236. Atlantic Highlands: Humanities Press.
516. Sir Karl Popper. Arch.Psychiatr.Nervenkr. 232, 1-3.
517. Biographical notes on Janos Szentágothai. Trends Neurosci. 5, 418-419.
518. How the self acts on the brain. Psychoneuroendocrinology, 7, 271-283.
519. Animal consciousness and human self-consciousness. Experientia, 38, 1384-1391.
520. Editor. Mind and brain: the many-faceted problems: selected readings from the Proceedings of the International Conferences on the Unity of the Sciences. Washington: Paragon House.
521. L'uomo e la scienza. In: La cultura strumento di ripresa della vita. pp.24-32. Milano: Jaca Books.

1983

522. The human mystery:a lifelong search for truth. Civitas, 38, 301-305.
523. The horizontal (tangential) fibres system of lamina I of the cerebral cortex. Acta Morphol.Hung. 31, 261-284.
524. Neurobiologia de la intencionalidad: la iniciacion des movimientos voluntarios. Campus, Revisita de la Universidad de Alicante, Primavera, 7-14.
525. Closing remarks. In: Towards the illimitable future in nervous system regeneration (ed. B. Haber, J.R. Perez Polo, G.A. Haslim & A.M.G. Stella), pp.397-405. New York: Alan R. Liss.
526. Das Zeitgefuhl. Naturw.Rdsch.Stuttg. 36, 427-432.
527. The mystery of personal existence. Scholar and Educator, 7, 5-18.
528. Calcium in long-term potentiation as a model for memory. Neuroscience, 10, 1071-1081.
529. The human brain: the challenge to science. IBRO News 11, 13-16.

1984

530. Physiological and pharmacological investigations on pain control. Schweiz.Mschr. Zahnmed. 94, 1004-1013.
531. (with D.N. Robinson) The wonder of being human: our brain, our mind. New York: Free Press; London: Collier Macmillan. (Translated into German and Italian.)
532. Review of Local cortical circuits by M. Abeles. Naturwissenschaften, 71, 54.

1985

533. In: Nobel Prize conversations with Sir John Eccles, Roger Sperry, Ilya Prigogine, Brian Josephson, with a commentary by N. Cousins. San Francisco: Saybrook.
534. Mental summation the timing of voluntary intentions by cortical activity. Behav.Brain Sci. 8, 542-543.

1986

535. Do mental events cause neural events analogously to the probability fields of quantum mechanics. Proc.R.Soc.Lond.B 227, 411-428.
536. L'uoma e il suo cervello. Fondmenti, 5, 133-149.
537. The investigations of N. Tsukahara on relocation of synapses on red nucleus neurones. Neurosci.Res. 3, 476-486.
538. Learning in the motor system. Prog.Brain Res. 64, 3-18
539. Chromatolysis of neurones after axon section. In: Recent achievements in restorative neurology 2: Progressive neuromuscular diseases (ed. M.R. Dimitrijevic, B.A. Kakulas & G. Vrbova), pp.318-331. Basel: S. Karger.
540. Chemical transmission and Dale's principle. Prog.Brain Res. 68, 3-13.
541. Reductionism and upward and downward causation. Biology Forum 79, 381-405.
542. Mechanisms of long-term memory. J.Physiol. Paris 81, 312-317.

1987

543. The effect of silent thinking on the cerebral cortex. In: The brain-mind problem: philosophical and neurophysiological approaches (ed. B. Gulyas), pp.31-60. Leuvin: Leuvin University Press.
544. Mechanisms of learning in complex neural systems. In: Handbook of Physiology, Sect.1. The nervous system, part 1, 137-167. Bethesda: American Physiol.Soc.
545. Neurologically effective nerve growths in the mammalian brain: recent work of Tsukahara and Kawaguchi. In: Clinical aspects of sensory motor integration (ed. A. Struppler & A. Weindl), pp.317-323. Berlin: Springer-Verlag.
546. Recollections on the early concepts of the synapse. Synapse, 1, 131-132.

1988

547. Mammalian systems for storing and retrieving memory. In: Cellular mechanisms of conditioning and behavioural plasticity (ed. C.D. Woody. D.L. Alkon & J.L. McGaugh), pp.289-302. New York: Plenum Press.

1989

548. The evolution of the brain: creation of the conscious self. London, New York: Routledge.

1990

549. A unitary hypothesis of mind-brain interaction in the cerebral cortex. Proc.R.Soc.Lond.B 240, 433-451.
550. Physics of brain-mind interactions. Behav.Brain Sci. 13, 662 .
551. Editor with L. Deecke & V.B. Mountcastle. From neuron to action: an appraisal of fundamental and clinical research. Berlin, New York: Springer-Verlag.
552. Editor with O. Creutzfeldt. The principles of design and operation of the brain. Vatican City: Pontificia Academia Scientiarum.
553. The mind-brain problem revisited. In: The principles of design and operation of the brain (ed. O. Creutzfeldt & J.C. Eccles), pp.549-572. Vatican City: Pontificia Academia Scientiarum.
554. Developing concepts of the synapses. J.Neurosci. 10, 3769-3781.
555. Evolution of consciousness. Proc.Natn.Acad.Sci. USA 89, 7320-7324.

1992

556. (with F. Beck) Quantum aspects of brain activity and the role of consciousness. Proc.Natn.Acad.Sci.USA 89, 11357-11361.
557. Keynote address. In: Rethinking neural networks: quantum fields and biological data. Part 6 of 13 video cassettes, in Ist Appalachian Conference on Neurodynamics (org. K.H. Pribram). Radford University Telecommunications Bureau.
558. The implications of nuclear war on the human being. In: The world-wide implications of nuclear war (ed. E. Etim & S. Stipcich), pp.163-169. 1st Internat. Sem.Nucl.War (1981). River Edge, NJ: World Scientific Publ.Corp.
559. A NATO policy of strengthening conventional weapons and forces. In: How to avoid a nuclear war (ed. E. Etim & S. Stipcich), pp.61-71. 2nd Internat.Sem.Nucl.War (1982). River Edge, NJ: World Scientific Publ.Corp.
560. Planning for peace. In: The technical basis for peace. (ed. S. Stipcich & W.S. Newman), pp.335-340. 3rd Internat.Sem.Nucl.War (1983). River Edge, NJ: World Scientific Publ.Corp.
561. Global cooperation. In: The nuclear winter and new defence systems: problems and perspectives. (ed. W.S. Newman & S. Stipcich), pp.468-471. 4th Internat.Sem.Nucl.War (1984). River Edge, NJ: World Scientific Publ. Corp.
562. Global freedom for scientific interchange. In: SDI, computer simulations, new proposals to stop the arms race (ed. W.S. Newman &Aamp; S. Stipcich), pp.355-357. 5th Internat.Sem. Nucl.War (1985). River Edge, NJ: World Scientific Publ. Corp.
563. East-West, North-South. In: The great projects for scientific collaboration east-west-north-south (ed. M.Dado), pp. 153-155. 7th Internat.Sem.Nucl.War (1987). River Edge, NJ: World Scientific Publ. Corp.
564. Features of the brain which are of importance in understanding the mode of operation of toxic substances and the radiation. In: From confrontation to co-operation (ed. M. Dardo), pp 154-165. 8th Internat.Sem.Nucl.War (1988). River Edge, NJ: World Scientific Publ. Corp
565. The closing session. In: The new emergencies (ed. M. Dardo & K. Goebel), p. 325. 9th Internat.Sem.Nucl.War (1989). River Edge, NJ: World Scientific Publ. Corp

1994

566. How the brain gives the experience of pain. Int.J.Tissue React. 16, 3-17.
567. How the self controls its brain. Berlin, New York: Springer-Verlag.

1998

568. (with F. Beck) Quantum processes in the brain: A scientific basis for consciousness. Cognitive Studies 5, 95-109.

John Moore was born in Lungling, China on 3 April 1941 and died in Canberra on 19 January 2013. 

He was an electrical engineer who spent most of his distinguished career at the University of Newcastle and the Australian National University following industrial experience and graduate education in Silicon Valley, California. 

He was a Fellow of the Institute of Electrical and Electronic Engineers, the Australian Academy of Science and the Australian Academy of Technological Sciences and Engineering, achieving all honours at a comparatively early age, and was recognised principally for his contributions to the field of control systems.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 25(1), 2014. It was written by Brian D. O. Anderson, Research School of Engineering, Australian National University.

Joan Mary (Jan) Anderson pioneered the investigation of the molecular organisation of the plant thylakoid membrane, making seminal discoveries that laid the foundations for the current understanding of photosynthesis. 

She grew up in Queenstown, New Zealand, obtaining a BSc and MSc at the University of Otago in Dunedin. After completing her PhD at the University of California, she embarked on a glittering career at the Commonwealth Scientific and Industrial Research Organisation (CSIRO) and then Australian National University (ANU) in Canberra. Not only a gifted experimentalist, Jan was a creative thinker, not afraid to put her insightful and prophetic hypotheses into the public domain. 

Her many notable achievements include establishing the details and the physiological significance of lateral heterogeneity in the distribution of the two photosystems between stacked and unstacked thylakoid membranes and the dynamic changes in the extent of stacking that occur in response to changes in the light environment. Her investigations brought her into collaboration with prominent researchers throughout the world. Recognised with many honours as a leading scientist in Australia, international recognition included Lifetime Achievement Award from the International Society of Photosynthesis Research, and Honorary Fellowships at Universities in the UK and USA.

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Supplementary material

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 30(1), 2019. It was written by Peter Horton, Wah Soon Chow and Christopher Barrett.

James W. Lance was a clinical neurologist who created the first university-based department of neurology in Australia. He championed academic enquiry and the scientific basis of clinical practice, and his research had two major themes, motor control and headache. 

After his doctoral studies on the pyramidal tract of the cat, he became a pioneer of the new field of motor control studied in human subjects, making seminal contributions on the control of muscle tone, reflexes and movement in healthy subjects and the pathophysiology of movement disorders in patients. At the same time he developed a clinical research program into the mechanisms and management of headache, in particular migraine. These studies evolved into parallel experiments in human subjects, cats and monkeys, probing the control of the cerebral circulation and the mechanisms underlying craniofacial pain, for which he received international acclaim in both fields. 

He received international and Australian honours and was the first practising clinician to be elected a fellow of the Australian Academy of Science. He is rightfully credited with leading the development of academic neurology in Australia and overseas.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 32(2), 2021. It was written by David Burke.

Written by David J. Collins, Gregory W. Simpson, David H. Solomon and Thomas H. Spurling.

Introduction

James Robert Price (who early in life became known as Jerry Price) was one of Australia’s leading chemists. After a distinguished career at the University of Adelaide and at Oxford University, where he worked with Professor Sir Robert Robinson, he returned to Australia following the Second World War with wife Joyce to join the then CSIR. He was to participate in a project initiated by the Division of Plant Industry, a search for medicinal drugs in the Australian flora, providing expertise in chemical studies; in time, he developed the project into the extraordinarily productive and coperative Australian Phytochemical Survey. He became Chief of the CSIRO Division of Organic Chemistry, then a member and eventually Chairman of the CSIRO Executive. In these roles he displayed great organisational and leadership skills. These were particularly needed during his time as Chairman when major changes in the structure of CSIRO were proposed by the government of the day. He was able to preserve the structural integrity and scientific focus of CSIRO during that period. He made major contributions to the discipline of chemistry in Australia particularly through his leadership and redirection of the Royal Australian Chemical Institute and his belief in the need for active interaction between Australian research institutes. He enjoyed an active retirement before suffering the effects of an unfortunate accident. He spent the last years of his life, supported by Joyce, in a local nursing home where he died on 8 March 1999.

Family background and early education

James Robert Price was born on 25 March 1912 at Kadina, a small town at the top of the Yorke Peninsula in South Australia. He was the eldest of three children reared by Edgar James Price (1875–1937) and Mary Katherine Price (née Hughes, 1883–1937); the other children were John (b. 1915) and Mary (b. 1917).

James Robert’s paternal grandfather, Benjamin James Price, after spending his early years in Kidderminster, England, migrated to Australia in 1857, lured not by the gold rush but by other opportunities offered by the new colony of South Australia. At the time of his marriage to Ellen Mary Carruthers in 1865, Benjamin Price was a Commission Agent in Adelaide; some time later they moved to Kapunda, a copper mining town, where their son Edgar James Price was born in 1875.

James Robert’s paternal grandmother, Ellen Mary Carruthers, was descended from a Scottish family that lived on the coast of Solway Firth, south of Annan in the shire of Dumfries. Some time in the 1840s Ellen’s father John Carruthers (1806–1887) migrated to South Australia where he established himself as a wine merchant in Adelaide. The Carruthers family connection is doubly strong because James Robert Price’s maternal grandfather, Henry Chauntrell Hughes, married Mary Catherine Carruthers, a cousin of Ellen Mary Carruthers.[1]

At the time of his marriage to Mary Hughes, Edgar Price was employed by the Savings Bank of South Australia in Adelaide; soon after that he was appointed as the first manager of the new Branch of that institution at Mount Gambier, and subsequently as manager of the Branch at Kadina, where his son James Robert Price was born. About two years later the Bank moved Edgar Price back to the managership at Mount Gambier, but when they asked him to move again, he decided to resign in order to establish his own accounting business; this was originally a partnership but he later operated on his own. Mount Gambier remained the family’s home town and it was here that at the age of about six James Robert Price began his education, first at a small Church of England school run by a Miss Warren, and then at Umpherston College, a Presbyterian school in Mount Gambier; Umpherston College was essentially a girls’ school, but included a minority of boys. In 1923 James Robert was awarded a Vansittart scholarship to St Peter’s College, an élite boys’ school in Adelaide. For the first two years he was a boarder in the preparatory part of that school. He then spent four years in the Senior School where he was originally a boarder, but when the economic depression hit Australia in the late 1920s Edgar Price was unable to afford the fees, and for the last year or so of his secondary education James Robert lived with his mother’s aunt Mrs Edith Helen Turner (née Carruthers) who was ‘very helpful to me but something of a dictator’.[2]

It was at St Peter’s College that Price was given the nickname ‘Jerry’ but he remained ‘Bob’ to his family – hence his choice, in due course, to be known as ‘Sir Robert’ rather than ‘Sir James’ – the name ‘Jerry’ came to be used generally and, to all those on first name terms, he was ‘Jerry’ for the rest of his life.

Upon completion of his Leaving Honours year at St Peter’s College in 1928, Jerry wanted to enrol in the Faculty of Science at the University of Adelaide. In a letter dated 12 November 1984 to Dr Rupert Best, who was collecting information for his book on the history of the Chemistry Department of the University,[3] Jerry recalled the circumstances of his entry into the University of Adelaide and the nature of his work there, both as an undergraduate and as a graduate student. Most of the following facts about Jerry’s work, study and research at the University of Adelaide have been extracted from that letter.

University of Adelaide

Jerry had looked forward to undertaking a university course in science in the usual way. However, because of the deepening economic depression, Edgar Price could not finance his son’s enrolment and the added expense of his boarding away from home. Hence, at his father’s urging, Jerry applied for a cadetship in the Chemistry Department of the University. He was interviewed by Professor A.K. Macbeth [4] who appointed him as a cadet at the beginning of 1929, initially on a wage of 10/- per week and later 25/- per week. This position gave him the right to enrol in such courses as Macbeth approved, without payment of fees. Because of their workload, cadets took four years rather than the usual three to complete a BSc degree. Jerry’s duties as a cadet included the preparation of solutions required for qualitative and quantitative chemical analyses in the first-year practical classes, but he also had some contact with third-year students doing analytical work under Dr W.T. Cooke. [5] In 1931 Macbeth transferred him to the job of Lecture Demonstrator, which entailed preparing and carrying out the experiments demonstrated by the Professor in lectures to the first-year students. This meant that he was now located in the Prince of Wales Building where he came into more contact with third-year and Honours (fourth-year) students. For an enthusiastic undergraduate, this regular contact with research students must have been both enlightening and stimulating. One of the people he met in this new environment was Gordon Kingsley (Bill) Hughes who, after completing his BSc degree in 1929, had become a Demonstrator in the Chemistry Department, and who in 1934 was appointed as Assistant Lecturer in the Chemistry Department of the University of Sydney. They became lifelong friends, and their close friendship was to have particular significance in what became the Australian Phytochemical Survey.

In 1933 Jerry began his research career as an Honours student under Macbeth’s supervision. In his letter to Best, he describes in some detail the research interests of Macbeth in the 1930s, and his own involvement as a research student. One interest that Macbeth brought with him from the University of Durham was the theoretical attempt to explain the rates of reaction of functional groups in unsaturated, particularly aromatic, compounds developed by Lapworth [6] in 1920 and called ‘induced alternate polarity’; another was the application of ultraviolet absorption spectroscopy to organic chemical problems. In Adelaide, Macbeth soon became actively involved in natural products chemistry, in part because his predecessor Professor E.H. Rennie had worked on the constituents of several Australian plants and had isolated two novel red pigments from the tubers of the insectivorous plant Drosera whittakeri that grows in the Adelaide Hills. Rennie had suggested that these pigments were probably naphthoquinone derivatives, but their structures remained unknown.[7] Here was a challenge, and one of the first projects that Macbeth gave to Jerry as an Honours student was to measure the ultraviolet absorption spectra of a number of known hydroxynaphthaquinones for comparison with the spectra of the Drosera pigments (4).[8] This enabled deduction of the structures of these pigments that Macbeth named droserone and Hydroxydroserone.[9] The structure of the latter was firmly established by synthesis by Winzor,[10] a junior and previously unproductive member of the staff whom Macbeth had stimulated into action.

Macbeth was further drawn into natural products chemistry by the fact that P.A. Berry and T.B. Swanson, two full-time employees of the manufacturing company A.M. Bickford and Sons,[11] came to the Chemistry Department as evening MSc students with the desire to work on components of the essential oil of Eucalyptus cneorifolia, a product of commercial interest to Bickfords. In a reflection on ‘scientific pedigree’ in his letter to Best, Jerry recalled: ‘I have little doubt that Macbeth’s preparedness to move into that field as presented to him by Berry and Swanson was influenced by his earlier contact with Read’. When Macbeth was a Senior Lecturer at the University of St Andrews, prior to his Readership at the University of Durham, the professor there was the terpene chemist John Read who, prior to that, had succeeded Robert Robinson in the Chair of Chemistry at the University of Sydney. While at Sydney, both Read and Robinson had collaborated with H.G. Smith, the terpene chemist who is regarded as the father of organic chemistry in Australia. In connection with the investigations of Berry and Swanson, Macbeth gave Jerry the job of making and characterising the 2,4-dinitrophenylhydrazone of the ketone (-)-4-isopropylcyclohex-2-en-1-one, one of the constituents of the essential oil of Eucalyptus cneorifolia. In the course of this work Jerry observed that the reagent 2,4-dinitrophenylhydrazine (which had only been developed in 1931) was broken down by alkali, and on his own initiative he began studying the products. He recalled:

Macbeth, rather than suggest that this was an unnecessary diversion from the main objective, encouraged me to pursue this study and further, when one of the products of the reaction was established as a 1,2,3- benztriazole [sic; 1-hydroxy-5-nitro-1H 1,2,3-benzotriazole (2)], he suggested an ultraviolet spectroscopic study of 1,2,3-benztriazoles, which was carried out.

Jerry graduated BSc (Hons, first class) in December 1934, and MSc in July 1935 with a thesis entitled ‘Properties of Nitro- phenylhydrazines and Absorption Spectra of Dimethyl-cyclohexanediones and 1:2:3-Benzotriazoles’. He was very industrious and efficient — he was a co-author of four papers (1–4) published in the Journal of the Chemical Society (London) in 1934/35, all of them having been received by the Editor between April and December 1934. He was co-author of three more papers (5–7) with Macbeth between 1935 and 1937. Jerry had high praise for his former supervisor: ‘Macbeth was a very good research supervisor – he presented research problems to those moving into research, he encouraged them to ‘do their own thing’ but he also kept a light rein on their movements’. The regard was obviously mutual – Jerry’s initiative, skill and industry were rewarded. In 1935, following Macbeth’s strong recommendation, Jerry was one of two Australians awarded an 1851 Exhibition Science Research Scholarship for that year. He accepted this to work as a DPhil student under Professor Sir Robert Robinson at Oxford.

In 1933, his Honours year, Jerry met his wife-to-be, Joyce Ethel Brooke, daughter of Roy Brooke and Myrtle Victoria Brooke (née Lackington). Joyce was in her first year as a science undergraduate at the University of Adelaide, and Jerry was her Demonstrator in the First Year Chemistry Laboratory: he escorted her to the Science Ball later that year. From about mid-1934 Jerry was President of the University Science Association and when he left for England in 1935 Joyce succeeded him as the first female President of that body. She graduated BSc (Hons, first class) in botany in 1936 and then worked at the Waite Agricultural Research Institute in Adelaide, doing research that gained her the degree of MSc in 1939, just before she set out for England to join Jerry.

Jerry’s ‘apprenticeship’ in research at the University of Adelaide was very successful, and set the course of his career: ‘my future career was the result of Macbeth’s interest in natural product chemistry’. On 5 September 1935 he boarded the Blue Funnel Line steamer ‘Nestor’ with a free passage to England, a privilege then accorded to 1851 Exhibition and Rhodes Scholarship holders by the Blue Funnel Line.

Oxford, the John Innes Horticultural Institution, and War

For two years Jerry lived at Magdalen College, Oxford while working at the Dyson Perrins Chemical Laboratory. His supervisor, Professor Sir Robert Robinson, was pre-eminent in the chemistry and biogenesis of plant natural products, and also in the synthesis of natural products.[12] One of Robinson’s interests at that time was the chemistry of plant pigments, including anthocyanins and their aglycones, the anthocyanidins. Jerry worked on the anthocyanidins in Bougainvillaea glabra and graduated DPhil in 1937 with a thesis entitled ‘Colouring Matter of Bougainvillaea glabra’.

Upon completion of the work for his DPhil at Oxford, and with Robinson’s recommendation, Jerry was appointed as Head of the Chemistry Section at the John Innes Horticultural Institution at Merton Park in South West London, near Wimbledon. Sir Robert Robinson had a close association with the Institution and collaborative research continued with Price’s appointment there. At the time of his appointment Jerry was the only chemist on the staff – his colleagues were all eminent geneticists, and all of them were Fellows of the Royal Society.

The research carried out by Jerry at the John Innes Horticultural Institution was on plant pigments – mainly on isolating the anthocyanins and investigating their role in the genetic variation of flower colour. Some of this work was done in collaboration with Sir Robert Robinson: of eleven papers Jerry published from the John Innes Horticultural Institution (12–22), six were jointly with Robinson. The collaborative research included the identity of the yellow pigment of Dahlia variabilis, undertaken to facilitate a study of the inheritance of flower colour in Dahlia species; also, a study of the yellow Papaver nudicaule (Iceland poppy) from which was isolated a nitrogenous diglucoside. Another collaborative study was on the orange-red pigment dunnione from Streptocarpus dunnii; dunnione was shown to be a furano-1,2- naphthoquinone – at that time a new class of natural product.

In 1939 Jerry was awarded a Rockefeller Scholarship, and with this he planned to go to the USA, after travelling first to Australia so that he and Joyce Brooke could get married. The outbreak of the Second World War in September 1939, however, enforced drastic changes, both professional and personal. Because of the war, Rockefeller (Travelling) Scholarships were cancelled, and Jerry decided to stay in the United Kingdom to help in whatever professional wartime service might be required. He asked Joyce to come to England. This was no easy matter, since much commercial shipping was being sunk by the German air and naval forces. Mrs Brooke insisted that her daughter Joyce should fly, and gave financial assistance to help with the additional expense. Joyce journeyed by train from Adelaide to Sydney via Melbourne to catch the Sunderland flying boat that left from Rose Bay, Sydney. The flight from Sydney to Poole in England usually took 10½ days. Joyce’s flight made overnight stops at Darwin, Sourabaya, Singapore, Rangoon, Calcutta, Karachi, Basra, Athens, Corfu (a replacement for the scheduled stop at Naples due to bad weather over the Appenines) and Marseilles; bad weather over the English Channel caused another stopover at St Nazaire before the flying boat reached Poole, 11½ days after leaving Sydney!

Jerry met Joyce at Poole after a separation of five years and they made plans to get married as soon as possible. On Tuesday 19 March 1940 they arrived in London and proceeded to Wimbledon, where Jerry was living because of its proximity to the Horticultural Institution at Merton Park. Temporary accommodation was arranged for Joyce nearby, and the Vicar of Wimbledon married them on Easter Saturday, 23 March 1940. In the rushed wartime wedding, away from home, there were no relatives present: the only attendants were Bill Hughes and his wife Jean, and Jerry’s former landlady Mrs Lacey. Hughes had recently arrived in England to spend a year at Oxford with Robinson. Joyce recalls that it was joked at the time that Bill Hughes was the best man and also gave her away, Jean Hughes was the congregation, and Mrs Lacey wore black and wept because Joyce could not have any family present. The wedding breakfast was a pie in Lyons Corner House! They had a two-week walking honeymoon in the country near Oxford, and on one evening were entertained to tea by Sir Robert and Lady Robinson.

Back at Wimbledon, the newlywed couple rented a flat on the top of a two-storey building near the railway line. This location was far from ideal – on moonlit nights the Germans bombed along railway lines. Rather than go to the huge air raid shelters, Jerry and Joyce slept in their flat, fully dressed, under a solid dining room table bought for £2.

While awaiting professional assignment in war service, Jerry served in the Home Guard (‘Dad’s Army’) and Joyce worked in first aid stations. In September 1941 Jerry was directed into the Chemical Inspection Department, Ministry of Supply, to supervise work in a group of five ICI factories in southwest Scotland in the manufacture of explosives and munitions. Initially they had six months in Ardrossan while Jerry familiarized himself with the production of munitions at Ardeer, the central ICI factory for the manufacture of explosives. The manager there was a Mr Lumsden, father of Harry C. Lumsden whom Jerry had met in his first year at Oxford. At that time Harry had taken Jerry to his home in Scotland for a holiday, so arising from this happy coincidence the Lumsdens became, as Lady Price recalls, ‘our family in Scotland’. After leaving Ardeer, Jerry became Chemist-in-Charge at the Powfoot Outstation, and later was in charge of the Dumfries area.[13] He and Joyce lived close by in the Royal Borough of Annan, and later in the rural area of Glen Stuart, both places only a bicycle ride from Powfoot. Life there was much safer than in frequently-bombed Wimbledon. Jerry supervised work on propellants and explosives, but no details are available as he left no written records of this period of his career.

During the Second World War Australia had to become self-sufficient in the supply of certain drugs. One of these was the anti-seasickness drug hyoscine, very important for naval operations. In late 1940, through the enterprise of Russell (later Sir Russell) Grimwade and his company Felton Grimwade and Duerdins, a part of Drug Houses of Australia, seven ounces of hyoscine were quickly extracted from 108 pounds of leaves of the Australian native tree Duboisia myoporoides picked near Grafton, New South Wales. Production was soon scaled up and hyoscine was produced during the war in quantities sufficient to supply British and American as well as Australian needs.[14] At the same time, CSIR became involved in a search for other drugs from Australian plants and in 1945, in anticipation of the end of the war, CSIR advertised a position of Research Officer for an organic chemist to work in the then Division of Industrial Chemistry on a survey of Australian native plants for sources of potentially useful alkaloids. Jerry Price was the successful applicant, and he and Joyce embarked on the first civilian passenger ship to make the trip between England and Australia since the war began; by coincidence, this was the ‘Nestor’, the ship on which Jerry had travelled to England in 1935. They embarked at Liverpool in early July 1945 with their two children born in Scotland – Margaret Ann (b. March 1944) and Donald Carruthers (b. May 1945). They arrived in Melbourne in the first week of September, ten years to the day since Jerry had left, having heard of the Japanese surrender during the voyage. Their third child, Janet Elizabeth, was born in Melbourne in October 1946.[15]

CSIR/CSIRO

The circumstances leading to the appointment of Jerry Price as a Research Officer in the CSIR Division of Industrial Chemistry have been described in some detail in a paper on what became known as the Australian Phytochemical Survey (72). The key points may be summarised as follows:

CSIR’s first involvement in Australia’s efforts to become self-sufficient in some key drugs during the Second World War was in mid-1940. Dr C. Barnard, Chief of the CSIR Division of Plant Industry, organised extensive field cultivation of some exotic drug plants: extraction of alkaloids and other active constituents from these plants was carried out by Dr H. Finnemore, Head of the Department of Pharmacy, University of Sydney. His limited resources meant that he was unable to cope with the ballooning demand and early in 1941 Professor R.D. (later Sir Douglas) Wright of the Department of Physiology, University of Melbourne, became involved. In his department some native Australian plants that were regarded as potentially interesting were extracted, and bioassays carried out by the pharmacologist Dr F.H. Shaw and his colleagues.

A very important development was Barnard’s appointment in June 1944 of Dr L.J. Webb as a ‘peripatetic botanist’ to boost the rate of collection of native plant species for chemical examination. Webb’s intimate knowledge of the flora of Queensland and his keen interest in poisonous plants and bush medicines,[16] together with his great drive and enthusiasm for the project, led him to provide important stimulation in communications with Jerry’s CSIR(O) group and collaborators from the Chemistry Departments of several Universities.[17]

Soon after Webb’s appointment it became apparent that there was a need within CSIR for an organic chemist who could give undivided attention to the isolation and characterisation of alkaloids and other secondary metabolites, and who could determine the structures of previously unknown compounds that were isolated. In September 1944, Barnard gained the enthusiastic support of Dr I.W. (later Sir Ian) Wark, Chief of the CSIR Division of Industrial Chemistry, for the appointment of a Research Officer for the Organic Chemistry Section of the Division. The position advertised was ‘for work on alkaloids from Australian native plants and trees’. The successful applicant, Jerry Price, was eminently qualified for the position; 16 of his 21 pre-1945 research publications were on the chemistry of plant extractives.

It is of interest to note that, while Jerry included a testimonial from Sir Robert Robinson in his letter of application, Guy Gresford, the Australian Scientific Research Liaison Officer in London, still wrote directly to Sir Robert for a reference. Sir Robert’s reply [18] was succinct:

Dear Gresford
In connexion with the appointment to the staff of C.S.I.R. I can strongly support J.R. Price who is an applicant.

The Minister in Charge of Scientific and Industrial Research, J.J. Dedman, approved Jerry’s appointment to CSIR on 15 March 1945 and he commenced duty on 24 September 1945.

Jerry’s initial scientific objectives were: to discover new sources of alkaloids already of value for medicinal purposes or as insecticides; to discover new alkaloids which may replace or supplement those already in use, and; to work out satisfactory methods of isolating and purifying the alkaloids found and to increase our systematic knowledge of the nature and chemistry of the alkaloids elaborated by plants.[19]

By the time Jerry arrived in Melbourne in 1945, Len Webb’s screening tests (some on freshly collected leaves and bark and some on herbarium specimens) had already identified a large number of alkaloid-containing plants. In October 1945, one month after taking up his position in CSIR, Jerry joined Webb at Innisfail, North Queensland, to collect bulk quantities of leaf, bark and wood of several selected species of rainforest trees belonging to the plant family Rutaceae for extraction and detailed chemical examination.

He very quickly impressed Dr Wark, who wrote, as part of a reclassification case on 18 February 1946:

Price is an excellent organic chemist and within a few weeks of his arrival in Melbourne was already obtaining results of importance in the alkaloid project. An 1851 Scholar, Price has a fine personality, is enthusiastic, and is a skilled experimentalist. He is quite capable of taking over the whole responsibility for the chemical side of the alkaloid investigation now being carried on in conjunction with the Division of Plant Industry and the Physiology Department of the University of Melbourne. It is recommended that he be reclassified as Senior Research Officer on 1/1/47, with a salary of £650 p.a., which is little enough for a man of his attainments who is almost 34 years of age.

It was already apparent to Jerry that to make significant impact in carrying out thorough chemical studies on the large body of Australian flora already known to give alkaloid-positive tests, many research chemists would be needed. He therefore sought help from staff in the Chemistry departments of the Australian universities. This was a remarkable achievement in university/CSIR(O) cooperation, brought about with a minimum of bureaucratic involvement. Probably one of the first university chemists whom he contacted was his good friend Bill Hughes at the Chemistry Department, University of Sydney. For some years, Hughes and his colleague Ern Ritchie (later Professor) had been studying anthocyanin pigments in Australian plants: they now enthusiastically took up the study of alkaloid-containing plants and many research students working for BSc Honours and MSc degrees cut their research teeth on the isolation and structure-determination of alkaloids. The other university chemist who collaborated with Jerry from the very beginning was F.N. (Norm) Lahey, a senior lecturer in the Chemistry Department of the University of Melbourne for the period 1943–1949 [20] before he took up a Research Professorship in Organic Chemistry at the University of Queensland.[21] Jerry’s collaboration with Lahey was facilitated and perhaps stimulated by the fact that the CSIR chemistry laboratories at Fishermans Bend were yet to be completed [22] and Price was provided with laboratory space and facilities in the Chemistry Department of the University of Melbourne through the cooperation and generosity of Professor E.J. Hartung. Although parts of the Fishermans Bend laboratories were completed and occupied much earlier, it was 1954 before Jerry and his group of phytochemists could move there.

Three of the six tree species collected by Price and Webb from the rainforest near Innisfail were Melicope (now Medicosma) fareana, Evodia (now Euodia) xanthoxyloides and Acronychia baueri (now Sarcomelicope simplicifolia): the detailed study of the constituent alkaloids of these three species was undertaken respectively by Price, Hughes and Lahey. All three species proved to be very rich in alkaloids, some of which were common to all three of the above Rutaceous plants. In 1948 Hughes, Lahey, Price and Webb published a note in Nature entitled ‘Alkaloids of the Australian Rutaceae’ (23). This first communication heralded what became a very large and very successful survey of Australian plants for alkaloids and other constituents of chemical and/or biological interest. The full details of Jerry’s work on Melicope fareana were set out in a series of papers in the Australian Journal of Scientific Research (24–27).[23] Parts II and V of this series were in collaboration with W.D. (Bill) Crow and Part III was by Crow alone. Crow was the first organic chemist appointed by CSIR to assist Jerry; he joined him in the laboratory at the University of Melbourne at the beginning of 1947 after having completed his BSc (Hons) degree at the University of Sydney with Hughes, who initiated him into the techniques of isolation and characterisation of alkaloids.[24]

The three rainforest tree species that featured in the 1948 note to Nature by Hughes, Lahey, Price and Webb gave the alkaloid survey a flying start. In addition to the five papers by Price and Crow on Melicope (now Medicosma) fareana, Lahey and his co-workers published four papers on the alkaloids of Acronychia baueri (now Sarcomelicope simplicifolia) in 1949–1950;[25] and from 1949 to 1952 Hughes and Ritchie and their collaborators published four papers on the alkaloids of Evodia (now Euodia) xanthoxyloides. [26] At the same time a number of other alkaloid-containing species were being examined by Jerry and his co-workers and by Hughes and Ritchie and their research students at the University of Sydney. The programme of chemical studies that Jerry initiated in 1945 had rapidly gathered momentum. After the early work on Melicope fareana, the Price CSIR(O)[27] group examined many other alkaloid-containing species while still working in their temporary accommodation at the University of Melbourne; those that featured in publications included Glycosmis pentaphylla, Pentaceras australis (now australe), Gyrocarpus americanus, Medicosma cunninghamii, Heliotropium europaeum; and Flindersia bourjotiana in collaboration with Hughes, Ritchie and Cannon [28] at the University of Sydney.

In 1946 the Chemistry Department of the University of Melbourne had a large increase in the numbers of students doing organic chemistry in the second year and it became necessary to divide the class: Professor Hartung obtained permission to appoint Jerry Price as a temporary part-time lecturer to give one lecture per week for the whole academic year; he was paid an honorarium of £100. [29] There were 200 students and the class was divided, Lahey taking one group and Jerry the other. This was a convenient arrangement with Jerry working in the Chemistry Department laboratories: it appears to have been operative only for the year 1946, but when Lahey moved to his new position at the University of Queensland in late 1949, Jerry became co-supervisor of some of his research students. One of these was Eva R. Klein (later Mrs Nelson); in her PhD thesis she acknowledged that ‘Part I of this work was carried out under the direction of Dr J.R. Price and Part II under the direction of Dr F.N. Lahey.[30] Part I of her thesis was ‘An Investigation of the Sulfur-Containing Alkaloid from the bark of Pentaceras australis Hook F.’ This project yielded three papers published in 1952 (31–33). Another PhD student supervised jointly by Price and Lahey was John A. Lamberton, who stated in his Acknowledgments that ‘This work was carried out under the supervision of Dr J.R. Price and, in part, Dr F.N. Lahey’. Lamberton worked on the alkaloids from Acronychia baueri and Medicosma cunninghammii [31] (28). After a postdoctoral year overseas, Lamberton was employed in 1951 by the CSIRO Division of Industrial Chemistry, to work under Dr Harold Hatt on plant waxes. In due course Jerry replaced Hatt as the leader of the Organic Chemistry Section. John Lamberton joined the phytochemical group in 1965 and became a major contributor to the expanding alkaloid program.[32] An MSc student who was supervised by Jerry was H.P. Haynes who worked on the alkaloids of Pentaceras australis.[33]

A search of the University of Melbourne staff files for the Chemistry Department [34] failed to reveal any correspondence with Professor Hartung formalising Jerry’s supervision of research students: the involvement of non-university personnel in such a role appears to have been less regulated than it is now. Perhaps the apparent lack of formalisation was a consequence of Jerry’s 1946 appointment as temporary part-time lecturer and his physical location in the Chemistry Department. The CSIR records show that Jerry was granted permission by CSIR to deliver a series of 25 lectures on Organic Chemistry to second-year students in 1946 but they do not indicate that he was appointed as a temporary part-time lecturer. In 1947 Jerry was invited by Professor Trikojus to deliver three lectures on plant pigments to advanced students of the Biochemistry Department, University of Melbourne. These are the only references to his involvement with the University in the CSIR(O) files.

The next organic chemist to join Jerry’s alkaloid group after Crow (1947) was L.J. Drummond (1948); then followed Dr N.V. (Noel) Riggs (1949) and Dr C.C.J. (Claude) Culvenor (1950). Culvenor replaced Drummond who moved to the Defence Laboratories, Salisbury, South Australia. Like John Lamberton, Claude Culvenor remained with CSIRO, going on to make a major contribution in the study of alkaloids, particularly those in plants poisonous to farm animals; in 1971 he moved to the CSIRO Division of Animal Health where this work continued. In 1951 Riggs took the Chair of Organic Chemistry at the University of New England and was replaced in the group by Dr Emery Gellert. All these chemists worked together with Jerry in the laboratory space provided by Professor Hartung in the Chemistry Department of the University of Melbourne, until they moved to the Fishermans Bend laboratories in 1954.

In 1952 Jerry, now a Principal Research Officer, gave the Liversidge Research Lecture at the 29th Meeting of ANZAAS, held at the University of Sydney (38). In this lecture he summarised the alkaloid work being carried out by the CSIRO and university chemists, but he also described other phytochemical studies being pursued in Australian universities.

In his introduction he referred to the early pioneering work on essential oils that had:

dominated Australian plant chemistry for the past fifty years. One result of this domination is a fine record of achievement in terpene chemistry, but another, inevitably, is that remarkably little has been accomplished with other classes of plant products. However, this situation is gradually changing today – plant chemistry in Australia rests on a broader basis than at any time in the past. In addition to essential oils, cellulose, lignin, tannins, waxes, colouring matters, triterpenes, steroids, alkaloids, coumarins, cyclitols and lignanes are being actively investigated. It is my intention to give you some idea of the kind of work being done and where it is leading.

Jerry played a key role in a conference held at the University of Melbourne in February 1947 on ‘Research into the Pharmacological and Other Chemical Constituents of Native Plants’. Six more such conferences were held (Melbourne, January 1949; Sydney, May 1951; Melbourne, August 1955; Adelaide, August 1958; Sydney, August 1962; and Melbourne, August 1965).

Jerry was awarded the DSc from the University of Adelaide in 1954 and was elected a Fellow of the Australian Academy of Science in 1959. Jerry was subsequently active on Academy business. He was a member of the National Committee for Chemistry from 1960 until 1966 and then Chairman of that committee 1969. From 1962 to 1968 he was a member of the Sectional Committee for Chemistry and Applied Chemistry (Chairman 1965–1966); a member of the Editorial Board of Records of the Australian Academy of Science 1965–1970; Chairman of the Publications Committee 1966–1970; and in 1969 a member of the Science and Industry Forum.

Of great significance for Jerry and for Australian phytochemistry was the decision of the International Union of Pure and Applied Chemistry (IUPAC) to hold a symposium on the Chemistry of Natural Products in Australia in 1960. The invitation to IUPAC was made in 1958 by Dr A.L.G. Rees, CSIRO Division of Industrial Chemistry, through the Australian Academy of Science. With Rees as Chairman of the Organizing Committee, Jerry responsible for the scientific programme and Sir Alexander (later Lord) Todd, Nobel Laureate, as President, the symposium was highly successful. Sessions were held successively in Melbourne, Canberra and Sydney (72, p. 347). Many eminent organic chemists from overseas participated, but particularly significant for Jerry was the presence of Sir Robert Robinson, who had retired from the Wayneflete Chair of Chemistry at Oxford in 1955 but was still very actively involved in chemical research. [35] This symposium put Australian chemistry, particularly the Phytochemical Survey, firmly on the international stage. Nine years later Jerry was chairman of the organising committee for two IUPAC conferences that were held at the University of Sydney in August 1969. These conferences attracted considerable media interest. A.L.G. Rees was at that time the President of IUPAC.

Jerry was promoted through the CSIRO research officer ranks until in March 1960 the then Officer-in-Charge of the Organic Chemistry Section, Dr Harold H. Hatt, informed the CSIRO Executive that he wished to resign from that position in order to devote his time to research. Hatt assumed the role of Head of the Sugar Research Group located at the University of Melbourne. Dr Wark recommended that Jerry be appointed to the vacant position. He wrote:

It is unnecessary to advertise this position. Throughout the world it would be regarded as certain that Dr Price would be offered it, and there is no prospect whatever of finding anybody more suitable for it. The Executive is well aware of Dr Price’s standing as an organic chemist, of his ability as an administrator and of the originality he has brought to bear with respect to the work of his group in the Organic Chemistry Section. I have no hesitation in stating that he has the capacity to lead a larger group with distinction.

CSIRO’s Chemical Research Laboratories [36] operated under the following terms of reference: to promote technical efficiency in established industries; to stimulate the establishment of new industries; to encourage the use of raw materials of Australian origin; to seek substitutes for imported materials; to find uses for by- products not utilised; and to study national problems to which its officers could contribute by virtue of their experience in other fields. [37]

These terms of reference are quite broad but the Annual Reports from that period indicate that individual scientists were given considerable freedom to develop separate projects within the overall strategy.

The Organic Chemistry Section was raised to Divisional status on 14 March 1961. Jerry was appointed as the first Chief of the new Division in which there were sixteen Research Officers supported by ten Experimental Officers. [38] The Division was thus only a fraction of the size of current CSIRO Divisions that now comprise 200–600 people. The five Divisions at Fishermans Bend were part of the Chemical Research Laboratories and until 1960 the Divisional reports were consolidated into the Annual Reports of the Chemical Research Laboratories. From 1961 separate introductory material was presented for each Division [39] and that from the Division of Organic Chemistry provides some insight into Jerry’s views on the purpose of the Division and how these influenced project selection.

In 1960 [40] projects were underway in the study of the chemistry of wool wax, sugarcane wax, long chain organic compounds (looking at insecticides and naturally occurring acetylenic compounds), brown coal tar constituents, the chemistry of stock poisons (largely pyrrolizidine alkaloids), the phytochemical survey of Australian plants, the chemistry of root exudates and the search for new uses for sugar. An important component of the work of the Division, and the Section before it, was done by the Microanalytical Unit, which provided a service to Australian universities and companies. Jerry often referred to the importance of the services that the Division offered to other organisations and of the interaction between all participants in Australian chemistry. The projects that had been referred to above had been in place with little change since 1951, apparently representing the continued work of the individual scientists. That is, it appears that scientists with particular research interests were appointed and that it was this research interest that dictated the choice of project, rather than projects being actively changed to reflect changes in strategic intent.

In his first Report as Chief, Jerry maintained that research in organic chemistry fell broadly into three categories: the investigation of natural products that provided much of the background information for the science, synthetic organic chemistry that was responsible for the tremendous output of the organic chemical industries, and physical organic chemistry. He stated that it was the first two of these that directly related to the utilisation of the country’s natural resources and to other national problems on which organic chemistry impinges, hence it was in those areas that emphasis was placed in planning the Division’s research. He indicated that much of the previous work in the Section had favoured natural products and that while this would continue he had initiated a reorientation that would broaden the synthetic activities of the Division. The principal feature of the new programme was the investigation of organic compounds of the metals of which Australia has large reserves, particularly aluminium, gold, zirconium, tungsten and titanium, with the objective of finding new uses for these metals. A second project closely integrated with the study of organometallic compounds was the study of organophosphorus chemistry.

In the 1962–63 Report [41] Jerry reported on the expansion of the Phytochemical Survey and the stock poison work. He again noted the importance of collaboration and reported that the Division was working with the CSIRO Division of Animal Health, the Western Australian Department of Agriculture, the Queensland Department of Agriculture and Stock, the Victorian Lands and Survey Department, Smith Kline and French Laboratories, the Cancer Chemotherapy National Service Centre, the US National Institutes of Health, the Population Council, the Queensland Department of Forests, the National Herbarium, the University of Sydney Department of Agricultural Microbiology, the University of Tasmania Department of Botany and Imperial Chemical Industries of Australia and New Zealand.

The 1963–64 Report [42] continues with a similar theme, again highlighting the service the Division provided to Australian chemistry.

The divisional projects that were underway when Jerry’s term as Chief concluded (at which time the Division of Organic Chemistry was merged with the Division of Physical Chemistry to form the Division of Applied Chemistry) illustrated how his views of the relevance of organic chemistry to Australia’s development had affected the Division’s work. He had terminated the work on the various waxes but had strengthened other areas. The projects reported in the 1964–65 Report [43] were synthetic organic chemistry aiming at the synthesis of new products of commercial value, organometallic chemistry, phytochemical investigations, toxic plant investigations and the recently introduced project on arthropod chemistry, principally considering the chemistry of insect and crustacean moulting hormones. At this time the impact of instrumental methods of chemical analysis was being felt and there was a corresponding decline in the reliance on microanalytical methods, coupled with a large decrease in the time required to determine chemical structures. The nature of organic chemistry had changed significantly away from bench chemical towards spectroscopic techniques and this was reflected in the acquisition by the Division of crystallographic, mass spectroscopic, NMR, and other major instrumentation. In the original establishment of the Division of Industrial Chemistry the spectroscopic work and the then relatively rare ultraviolet and infrared spectrophotometers were allocated to the Chemical Physics Section (later Division). By the mid-1960s it was clear that such expertise needed to be closely integrated with the organic chemists and Jerry recognised that scientists specialising in the application of the spectroscopic techniques were needed, rather than scientists concentrating on designing and developing new instruments.

Jerry was promoted to Chief Grade III on 1 July 1963 and wrote to the Chairman of CSIRO, Sir Frederick White, in characteristic style:

Dear Sir Frederick
I thank you very much indeed for your letter of 23rd July informing me of my reclassification to Chief Grade III. I appreciate this very much indeed though I must confess I don’t think it was warranted!

Royal Australian Chemical Institute (RACI)

Jerry felt strongly about the importance of professional relations between chemists in Australia, seeing effective interactions as critical to the success of the discipline. He was President of the Victorian Branch of the RACI in 1959 and Federal President from 1962 to 1964. He was Associate Editor of Institute publications (1949–1953), a member of the Editorial Board (1954–1955) and Editor of both Proceedings of the Royal Australian Chemical Institute and Reviews of Pure and Applied Chemistry (1956–1958).

Jerry was keen to promote the publication of Australian chemistry in the Australian Journal of Chemistry and to boost the image of the journal overseas. Jerry noticed that Australian publications appeared in Chemical Abstracts only after considerable delay. Enquiry revealed that Chemical Abstracts at that time preferred to have abstracting done in the country of publication, and that for Australia, this was in the hands of a person at CSIRO Head Office. To the abstractor’s relief, Jerry offered to take it over, arranging a group of people to help. Claude Culvenor, who had abstracted for British Abstracts, took this role for a period.

To understand the abstracting load, it must be remembered that in the 1950s and before, Chemical Abstracts wanted every chemical detail recorded in the abstract so that work was repeatable from the abstract. In organic chemistry each abstract required close reading and could be quite lengthy. By the late 1950s Chemical Abstracts recognised that this was too ambitious (also too costly, for publication and to purchasers) and abstracts began to be merely a summary of what was done. From then on, abstracting was done ‘in-house’ at Chemical Abstracts.[44]

In an article published in Proceedings of the Royal Australian Chemical Institute in September 1967 entitled ‘The Organization of Chemists and Chemistry’ (61), Jerry argued that Australian chemists must follow the example of chemists in the UK and unite their organisations rather than fragment them into special-interest groups. He described as regrettable the formation of the Australian Institute of Food Science and Technology and the Australian Oil and Colour Chemists Association, arguing that the best way forward was to adopt an organisational framework that integrated regional loyalties and interests with scientific and technological activities.

As mentioned above, Jerry was President of the Institute for an unprecedented two years from November 1962 to November 1964. When elected President he was already Chief of the Division of Organic Chemistry and in that position was on an overseas visit to: IUPAC Symposium on Pharmaceutical Chemistry in Florence (Italy); Dr J.S. Anderson, National Chemical Laboratory; Chemistry School, Cambridge University; ICI Pharmaceuticals Division, Research Laboratories; Department of Chemistry, University of Manchester; Boots Ltd, Research Department; Department of Chemistry, University of Nottingham; National Institute for Medical Research, Mill Hill; Smith, Kline and French Laboratories, Welwyn; School of Pharmacy, University of London (UK); International Symposium on Chemical Plant Taxonomy (France); Professor Kjaer, Copenhagen University (Denmark); Stockholm University, Professors Erdtman and Sandberg, Professor Sorenson, Trondheim University (Sweden); Division of Pure Chemistry, National Research Council, Ottawa (Canada); Dr Warren Nelson, Population Council, New York; National Institutes of Health, Bethesda, Maryland; Smith, Kline and French Laboratories, Philadelphia; Department of Chemistry, UCLA, Los Angeles; Department of Chemistry, University of Hawaii (USA). He left Melbourne on 13 September and returned on 21 November. The extensive list of contacts on this visit is an indication of how widely Jerry interacted with the international chemical community.

During this time he was also able to visit the headquarters of the Royal Institute of Chemistry, the Canadian Institute of Chemistry and the American Chemical Society, although none of these visits was mentioned on the official itinerary. On his return he wrote what was to be a very influential article for Proceedings of the Royal Australian Chemical Institute entitled ‘Home Thoughts from Abroad’ (53), in which he set out his views of the role and purpose of the RACI.

Jerry was very impressed with the structure of the Canadian Institute of Chemistry, particularly the disciplinary- based, nation-wide Divisions and the existence of a small Board separate from the larger Council. At that time the RACI did have an Australia-wide Cereal Chemistry Group and a newly-formed Polymer Group and Jerry strongly encouraged the formation of other divisions. These were the forerunners of the current Divisions of the RACI. He also floated the possibility of the Branches (state-based) encouraging more independent Sections. At that time there were Sections at Newcastle, Geelong and Canberra. He thought that there should definitely be Sections at the Latrobe Valley, Launceston, Wollongong, Ballarat, Port Pirie, Armidale and Bendigo. With some greater effort there could also be Sections at Broken Hill, Mount Isa, Kandos, Mackay and Townsville.

By 2003 there was in fact a Branch in Canberra and also in the Northern Territory, and Sections at Geelong, Gippsland and Ballarat/Bendigo in Victoria; New England, Newcastle, Northern Rivers, Riverina Murray, Western Sydney and Wollongong in New South Wales; and North Queensland, Central Queensland and Darling Downs in Queensland, even though centralisation of the chemical industry in Australia has prevented strong regional growth of the RACI of the type that Jerry may have envisaged. The RACI now has a highly successful Divisional structure, although it is still searching for an ideal operational model. In 2000 the RACI went to a new structure of the type suggested by Jerry, consisting of a Board to manage its business affairs, separate from an Assembly that sets policy direction and implementation.

The RACI awarded Jerry the H.G. Smith Medal (awarded in recognition of contribution to the field of organic chemistry) in 1956 and the Leighton Medal (its highest honour) in 1969.

Chairman of CSIRO

Jerry was appointed as a member of the CSIRO Executive from 27 January 1966 and attended his first meeting on 3 February 1966. He replaced another organic chemist, Professor G.M. Badger. His salary increased marginally from $12,108 to $13,200!

John Shelton recalls: [45]

In retrospect, Price’s appointment can be seen as the key step in Sir Frederick White’s objective of better preparing CSIRO to meet the political and bureaucratic assaults which he predicted would come. It was this apprehension that had prompted him to move to Canberra, to restructure Head Office and to replace retiring members of the Executive with younger and innovative policy makers; Ives, as a member, Lewis Lewis as associate member. It was the addition of Price, having the cachet of a former Chief, which gave the impetus to change, which Ives and Lewis could not themselves have achieved. Added to that was Price’s willingness to support changes that would benefit CSIRO, even though not popular with some chiefs – particularly a small but vocal and influential group who regarded CSIRO as a collection of ‘autonomous research institutes’. This group had already been suspicious of White’s restructuring of Head Office, following the departure of Mr. G.B. Gresford. In place of the single secretary as leader of Head Office, White created three secretaries, Administrative (L.G. Wilson), Agricultural & Biological Sciences (A.F. Gurnett Smith), and Industrial and Physical Sciences (J. P. Shelton).

On joining the Executive, Price made it widely known that he intended to make up his mind on what his role on the Executive should be after seeing from the inside how the Executive functioned. This was the first indication that Price was not a supporter of the ‘autonomous’ concept, and as it turned out, he would be largely responsible for the Executive becoming increasingly involved in what went on inside Divisions, especially in setting priorities and allocating resources at programme level. This came about first by the adoption of programme budgeting, and then through a greatly increased Executive involvement in detailed reviews of Divisions, such reviews having been in the past made only when a Chief had retired. However, that was to come later. Price’s first initiative on the Executive led to the Divisions of Organic Chemistry and Physical Chemistry being merged into a new Division of Applied Chemistry. He reasoned that the existing titles gave the impression that their role was to advance knowledge respectively in organic and physical chemistry. That, Price said, was a function of Universities, and distinct from the role of CSIRO in which these branches of chemistry were applied and, if necessary, further researched to provide solutions to industrial, economic, and in present day parlance, environmental problems, in terms of the Act, and of their respective divisional terms of reference. CSIRO was not, and should not seem to look like, a place where research was done for its own sake. CSIRO required both quality of research and relevance.

Price then took up his examination of Executive and Head Office procedures. He started information gathering with Shelton to assist but within a short time, Ives and Lewis had joined in what became a working party aimed at exploring and developing proposals for improvements in procedures in order to strengthen the effectiveness of the Executive and to prepare CSIRO to face the expected bids for external control of CSIRO. The key to this was formulated as follows – How can the Executive allocate funds and staff requested by a chief for growth without knowing, first, how much of current resources are going into that activity already in that and other divisions; second, how sure can the Executive be that the need for growth could not be better met, not by new funding, but by closing down unproductive or lower priority activities.

As it happened, the Finance group in Head Office had some time earlier appointed Nicholas Clarke to examine and report on the possible merit of CSIRO adopting the latest American management tool – programme budgeting. Clarke’s recommendation that this be adopted was turned down, as the finance group did not see how it could help them in their role, which was to use the Treasury headings of salaries, overtime, travel, equipment and so on. Clarke left. Shelton fed Clarke’s report into the working party, as the way to provide the Executive with the information it needed, as defined above. It was eventually adopted, initially as an information presentation to the Executives of the Industrial and Physical Sciences Divisions, then later to all Divisions and after Price became chairman, as the budget format.

It was a key change and enabled the Executive to be better informed, to allocate funds according to its priorities, and to be able to counter, in detail, the frequently raised canard that CSIRO was doing too much basic research. Programme budgeting identified each activity with a relevant industrial and economic problem which, if it could not be solved by applying existing knowledge, was being tackled by ‘basic’ research to seek new information that could lead to a solution, as it so often successfully achieved.

Professor David Solomon FAA, Foundation Chief of the CSIRO Division of Applied Organic Chemistry (1974 to 1987) and its successor the Division of Chemicals and Polymers (1988 to 1989), recalls that after Jerry had joined the executive of CSIRO he maintained an active interest in the business of the Division and, indeed, visited the Fishermans Bend Laboratories on one occasion to propose, after typically brief pleasantries, that Solomon attend a meeting at Thredbo that was to discuss the forgery threat to Australia’s currency. Professor Solomon attended the meeting and initiated the project that eventually, under Solomon’s leadership, produced the Australian polymer banknote technology. [46]

Jerry was appointed Chairman of the CSIRO Executive from 26 May 1970. The then Chairman, Sir Frederick White, wrote a personal letter to the Minister for Education and Science, Nigel Bowen, on 16 March 1970:

My dear Minister

I promised to let you know Dr Price’s plans for coming to Canberra. Dr Price will be renting a flat in Canberra in April and intends to spend most of his time here. This arrangement will persist until a house is available for him. He hopes this will be towards the end of the year and he will then bring his wife to Canberra.

Our building on Mt Ainslie is due to be completed in November and will, I think, be handed over to us not long after that. Plans are already afoot to move the staff from Melbourne to Canberra towards the latter part of the year.

An insight into Jerry’s integrity and modesty can be found in the discussion of his travelling allowance during this interim period in Canberra. He intended to commute to Canberra from Melbourne, spending the weekends in Melbourne. The maximum that he was entitled to was $105.00 per week if he chose the usual ‘per diem’ rates. He chose instead a modified package of $37.60 that included the actual cost of his flat and a special rate of $25.00 per week allowance.

One of Jerry’s first tasks as Chairman was to clarify the roles of the CSIRO Executive, the Head Office staff and the Chiefs. This was done for two reasons:

First, because I believe that the definition of these functions would be a worthwhile exercise in itself, but secondly, because it seems to me that we can’t usefully examine the relations between these three groups until we are reasonably clear as to their respective functions.

He studied the Science and Industry Research Act that governs CSIRO and came to the conclusion that:

It is therefore clear that the Executive, subject to the approval of the Minister, bears the responsibility for determining the policy of the Organization, for determining priorities and for allocating funds as it thinks best. But it goes a little further than this. In addition to determining policy or policies it is the responsibility of the Executive to see that such policies are implemented.

He was keen to develop a system of ‘cabinet solidarity’ with respect to Executive decisions and rejected the proposal of the Chiefs that individual members of the Executive should take line management responsibilities for particular activities such as finance and human resources. The consequence of this thinking was that the Executive had to rely on the Head Office staff to initiate the implementation of Executive decisions.

Jerry stated the role of the Chiefs in unambiguous terms:

While the Executive does call on you for advice and assistance in meeting its responsibilities, it also relies on you for many other things. You have the immediate responsibility for the first and most important function of the Organization laid down in the Act: ‘… the carrying out of scientific researches and investigations in connexion with or for the promotion of primary or secondary industry in the Commonwealth…’

In retrospect, Jerry was struggling with a management structure that did not separate the role of the Chairman from that of Chief Executive. This was not resolved until the appointment of a Board with an independent Chairman by the Hawke Government in 1986.

The election of the Whitlam Government on 2 December 1972 was the start of the most turbulent time in Jerry’s career. An insight into just how much he was affected by some of the decisions of that government’s ministers can be found in a speech that he gave to Melbourne staff on 3 July 1975, three and a half weeks after the Government announced that the Mineral Research activities of CSIRO were to be transferred to the Department of Minerals and Energy.

He started by saying: ‘This has been a period of considerable difficulty for all of us, and it is imperative that we now pause and look closely at the situation’. After congratulating the staff on their patience and restraint he continued:

it is necessary to go back about two years, CSIRO fully recognises its responsibility to do research to meet national needs and therefore to be responsive to Government policies. After the new Government came into office we immediately started thinking about our role in relation to the new Ministries that had been established. … As part of this communication exercise we drafted a letter to Mr Connor that Mr Morrison [47] signed on 5 April 1973 advising him of our research programme on solar energy and suggesting that discussions take place at officer level. Then on the 3 July that same year a review of our Minerals Research Programme went from Mr Morrison to Mr Connor. Mr Morrison invited Mr Connor to establish direct contact with CSIRO if he had any questions about the review, or if he wanted further information.

Unfortunately, no reply was received from Mr Connor to either of these letters.

Then in December 1973 Mr Connor made a statement in Parliament about the Government’s intention to undertake a crash programme on coal hydrogenation. Mr Morrison again wrote to Mr Connor telling him CSIRO was prepared to help in this programme and he asked Mr Connor to let him know details of the proposed crash programme.

Again, Mr Connor did not reply.

Jerry reminded his audience of a press statement dated 11 September 1974 in which the two Ministers had agreed that CSIRO would pursue research in many areas that would bear on the utilisation of solar energy and that the Department of Minerals and Energy would take over the development phase of those CSIRO results that were approaching practical realisation.

Many other attempts were made to work with the Department of Minerals and Energy but, according to Jerry, they were all ignored until 13 May 1975 when Mr Connor replied that at least the responsibility for liquid fuels from coal had been assumed by the recently formed Coal Conversion Sub-committee of the Coal Research Advisory Committee.

Jerry then gave a detailed account of developments after Thursday 5 June 1975:

Now to more recent developments. You probably all remember that on Thursday 5th June the Prime Minister issued a press statement outlining new Ministerial responsibilities. One sentence of his statement said, and I quote: ‘The Department of Minerals and Energy will take over responsibility for the Minerals Research Laboratories and the Solar Energy Studies Unit.’

... but let me reiterate my attitude, the attitude of the Executive and I believe the attitude of the Organization as a whole. This is that we object most strongly to the manner in which this decision was made and announced – without prior consultation with the former Minister for Science, Mr Morrison, his successor Mr Cameron, with the Organization, with the recently established ASTEC – the Australian Science and Technology Council – or with industry. We also object to the disregard of the practical requirements for conducting effective government-based scientific research in this country. We do not question the Government’s right to so order the affairs of CSIRO – that is its right – but such ordering should be brought about with the full understanding of all the factors involved and I believe we have preserved our credibility as responsible scientists and administrators by using every proper means to express our point of view to the Government. We have explained in very clear terms that the ad hoc dismemberment of CSIRO in this way could be disastrous to Australia’s scientific output for years to come.

Lady Price recalls that before Jerry went to give that speech he told her to expect that he would lose his job over his stand.

History records that the Government, already under pressure from adverse reactions to many of its decisions, and with the urging of the Acting Minister of Science and Consumer Affairs, Dr Moss Cass, finally rescinded the 5 July administrative orders. Jerry had thus very effectively preserved the integrity of CSIRO and enhanced his reputation both inside and outside the organisation.

Jerry retired on his 65th birthday, 24 March 1977. The Prime Minister, Malcolm Fraser, wrote:

I thought I should write to you on the occasion of your retirement as Chairman of the Executive of CSIRO.

My two periods as Minister of Education and Science enable me to write to you with first hand knowledge. I am well aware of the contribution you have made, successively, as an individual scientist, Chief of Division, member of the Executive and finally as Chairman of the Executive. Your appointment as Chairman designate was of course made while I was Minister responsible for CSIRO.

May I, as Prime Minister and personally, express my gratitude and appreciation for the service which you have rendered the Government and the nation, in your various capacities – but more particularly as Chairman of the Executive. The very high world wide standing of CSIRO (and CSIR before it) is due in no small part to the quality of the leadership over the years of its existence.

However I believe it is fair to say that your period as Chairman has occurred at a time when contemporary circumstances have never been more challenging. Throughout your service, your sense of dedication and loyalty, and your integrity, have been manifest for all to witness.

In conclusion, may I once again both thank and congratulate you for a job well done. I would like to express my very best wishes for a long and happy retirement.

Yours sincerely
(Malcolm Fraser)

Jerry was awarded KBE in 1976 in recognition of his services to science and government. He was made an honorary member of the Royal Society of New South Wales in 1977.

Retirement

Retirement afforded Jerry the opportunity to spend much more time with his grandchildren and to enjoy his garden. Jerry was also a great support to Lady Price in her many activities, as Lady Price was to him. When the Prices arrived in Melbourne in 1945, Joyce was aware of the great shortage of science teachers in secondary schools in Victoria. She suggested to Jerry that she embark on a teaching career but Jerry was not comfortable with his wife working. She abandoned that idea and went on to devote her time and considerable intellect and organising ability to the Girl Guides.

She rose to be the Victorian State Commissioner for Guides, the Chief Commissioner of the Girl Guides Association of Australia, 1968–1973, and the Chairman (for two terms) of the World Committee of the World Association of Girl Guides and Girl Scouts (1975–1981). She had been a member of that committee from 1972. Lady Price was made a Life Vice-President of the Girl Guides Association of Australia in 1984 and was honoured to speak at the funeral service for Lady Baden Powell.

Lady Price was in England on Girl Guide business in 1974 when Jerry rang her to say that he was likely to be dismissed because of the stand he was taking against the Government (see above). He reassured her that he would be able to return to a Research Scientist position!

In retirement Jerry served as a director of Humes [48] for seven years. He accepted the position in the hope that he could persuade the company to use Australian R&D to develop new technology rather than buying technology from overseas. While he did not succeed in that aim, he found working with business people intensely interesting and confided to Lady Price that he came across attitudes and processes that he had not experienced in his long career in public service science.

He was a member of the Monash University Council and on the Clunies Ross Foundation. In July 1986, Jerry was attending a meeting of the Clunies Ross Foundation at Clunies Ross House in Melbourne. He had parked his car on Royal Parade but was informed that a parking place had been reserved in the underground parking area at Clunies Ross House. Upon leaving the building via the back lane he was struck by a delivery truck. As a result of that accident Jerry suffered permanent lung damage and some brain damage. He withdrew from active public life after that.

After a period in the Shoreham Nursing Home Jerry died on 8 March 1999. A Memorial Service was held to commemorate his life on Friday 16 April 1999 at the Monash University Chapel where friends and colleagues were invited to speak and to contribute with others to a booklet commemorating Jerry’s life that was presented to Lady Price on the day on the service.[49]

In 1990 the CSIRO Division of Chemicals and Polymers (later CSIRO Molecular Science) instituted a named lecture series in tribute to Jerry’s influence on organic chemistry in CSIRO. There have been nine Sir Robert Price Lecturers to date, Mr Rod Rickards of the Australian National University in 1990, Dr Dan Kleier of DuPont Agricultural Products in 1991, Nobel Laureate Sir John Cornforth FRS of the University of Sussex in 1992, Professor Emmanuel Vogel of the University of Cologne in 1993, Associate Professor (now Professor) Max Crossley of the University of Sydney in 1994, Professor Paul Knochel of University of Marburg in 1996, Professor Steven Ley FRS of the University of Cambridge in 1998, Professor Tony Barrett FRS of the Imperial College of Science, Technology and Medicine in 2000 and Professor David Solomon FAA of the University of Melbourne in 2001. These lectures bring industry, CSIRO and university scientists together in a way that Jerry approved.

Sir Robert Price was a great organiser and project developer with the ability to make wide and useful contacts with influential people in associated fields. He made a significant contribution to the growth and development of chemistry in Australia, and to the development of public sector research. He was a great organic chemist and a great man.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 15(1), 2004. It was written by:

• David J. Collins, Honorary Senior Research Fellow, School of Chemistry, Monash University; Senior Fellow, Department of History and Philosophy of Science, University of Melbourne.
• Gregory W. Simpson, Deputy Chief, CSIRO Molecular Science.
• David H. Solomon, Professorial Fellow, Department of Chemical and Biomolecular Engineering, University of Melbourne.
• Thomas H. Spurling, Director, Industrial Research Institute Swinburne, Swinburne University of Technology.

Acknowledgments

The authors would like to thank Lady Price and the Price family for their encouragement and helpful comments and for making available much of the family historical material. We would also like to thank Dr Claude Culvenor and Mr John Shelton for helpful comments and suggestions on the manuscript. We thank Mr Rob Birtles of the CSIRO Records and Archives Strategies Group, Mr Michael Piggot, Archivist of the University of Melbourne, and staff of the Baillieu Library, University of Melbourne, for access to files and theses, and Mr Gavan McCarthy of the Australian Science and Technology Heritage Centre for access to archival material. The title of Jerry’s MSc thesis was kindly obtained by Dr George E. Gream. We thank Ms Carolyn Larsen of the Ian Wark Library, CSIRO, for assistance with historical and bibliographic material.

Notes and references

1. Information kindly supplied by Lady Price CMG, OBE; some details extracted from a handwritten document ‘My Family History’, by Catherine Joyce Price, granddaughter of Sir Robert Price. Various personal details outlined below about Price’s life have been drawn from documents made available by Lady Price.
2. J.R. Price, handwritten memoir in the possession of Lady Price.
3. Best, R.J., Discoveries by Chemists: A History of the Chemistry Department of the University of Adelaide, 1885–1984 (Adelaide, 1987). Parts of Sir Robert Price’s letter to Dr Rupert Best are quoted on pp. 85, 86. A copy of this letter was made available by Lady Price.
4. Alexander Killen Macbeth was appointed in March 1928 as successor to E.H. Rennie in the Angas Chair of Chemistry at the University of Adelaide: Best op. cit. (n. 3), pp. 75,78.
5. Dr William Ternent Cooke, who graduated from the University of Adelaide in 1900, won an 1851 Exhibition Scholarship which he took up in 1902 at University College, London to work with Sir William Ramsay. In 1906 Cooke was appointed as the first full- time Lecturer in the Chemistry Department, University of Adelaide [Best, op. cit. (n. 3), pp. 28–30)].
6. Lapworth, A., ‘A Theoretical Derivation of the Principle of Induced Alternate Polarities’, J. Chem. Soc. 121 (1922), 416–427.
7. Rennie, E.H., ‘The Colouring Matter of Drosera whittakeri’, J. Chem. Soc., 51 (1887), pp. 371–377; idem, ‘Notes on the Colouring Matter of Drosera whittakeri’, Trans. Proc. & Report of the Phil. Soc. Adelaide, 10 (1888), pp. 72–73; ‘On the Colouring Matter of Drosera whittakeri’, Report Second Meeting Australasian Assoc. Adv. Sci., 2 (1890), pp. 398–399; idem, ‘The Colouring Matters of Drosera whittakeri’, J. Chem. Soc., 63 (1893), pp. 1083–1089.
8. Numbers in brackets refer throughout to papers in the Bibliography below.
9. Macbeth, A.K., and Winzor, F.L., ‘The Colouring Matter of Drosera whittakeri. Part II’, J. Chem. Soc. (1935), pp. 334–336.
10. Winzor, F.L., ‘The Colouring Matter of Drosera whittakeri. Part III. The Synthesis of Hydroxydroserone’, J. Chem. Soc. (1935), pp. 336–8.
11. A.M. Bickford and Sons was a chemical drug manufacturing company in Adelaide that produced, inter alia, eucalyptus oil under the trade-mark ‘Our Jack Brand’; D. Shiel, Eucalyptus, Essence of Australia: The Story of the Eucalyptus Oil Industry – and of the ‘Eucy’ Men, and their Contribution to the Australian Bush Tradition, Queensberry Hill Press, 1985, pp. 204, 212: see also J.R. Poynter, Russell Grimwade, (Melbourne, 1967), pp. 31–33, 162. In 1930, A.M. Bickford and Sons amalgamated with a number of other companies to form Drug Houses of Australia Ltd. Felton, Grimwade and Bickford Pty Ltd: A Brief Chronological History, www.fgb.com.au.
12. Lord Todd and J.W. Cornforth, ‘Robert Robinson, 13 September 1886–8 February 1975’, Biographical Memoirs of Fellows of the Royal Society, 22 1976, pp. 415–527.
13. J.R.Price, notes on his early career in his 1946 application for Associate Membership of the Royal Australian Chemical Institute (RACI), The Records of the RACI, listed by Bill King, Gavan McCarthy and John Spink; Membership Files, Series 1, 1918–1956, Box 8; held at the Australian Science and Technology Heritage Centre, University of Melbourne.
14. Grimwade, J.F.T., A Short History of Drug Houses Australia Ltd to 1968 (Melbourne, 1974).
15. Margaret Ann Devlin née Price was trained as a physical education and mathematics teacher and is now the Deputy Principal of Strathcona Baptist Girls Grammar School, Melbourne; Dr Donald Carruthers Price is a physicist and is Senior Science Fellow with CSIRO Telecommunications and Industrial Physics, Sydney; Janet Elizabeth Price studied Russian and German at ANU and works with refugees.
16. In 1948 Webb published an important document that became very useful to participants in the Australian Phytochemical Survey; L.J. Webb, Guide to the Medicinal and Poisonous Plants of Queensland, CSIR Bulletin No. 232, 1–202, Melbourne, 1948.
17. Over a period of about 14 years (1949–1963) L. J. Webb wrote a series of confidential Phytochemical Newsletters, copies of which were distributed to all CSIRO and University personnel participating in the Australian Phytochemical Survey. Two photocopy sets of these have been bound by David Collins; one is held by him and one by L.J. Webb.
18. CSIRO Archives, Archive Box PH/PRI/1; the contents of this file were used extensively as the source of information about Price’s career in CSIRO.
19. Report of the CSIR Chemical Research Laboratories, 1945.
20. J. Radford, The Chemistry Department of the University of Melbourne: Its Contribution to Australian Science 1854–1959 (Melbourne, 1978), pp. 201, 227, 228.
21. B. Chiswell, A Diamond Period: A Brief History of the Chemistry Department of the University of Queensland 1910–1985 (Brisbane, 1986), p. 45.
22. I.W. Wark, ‘The CSIRO Division of Industrial Chemistry 1940–1952’, Records of the Australian Academy of Science, 4 (1979), 7–41. The building of the CSIR chemical laboratories at Fishermans Bend was authorized in 1946; work began in 1948 but was not completed until 1954.
23. After volume 5 (1952) the Australian Journal of Scientific Research, Series A Physical Sciences, was differentiated into the Australian Journal of Chemistry and the Australian Journal of Physics, each beginning with Volume 6 (1953). Publication of the Australian Journal of Scientific Research, Series B Biological Sciences also ceased in 1952 and was replaced by the Australian Journal of Botany and the Australian Journal of Zoology, each beginning with volume 1, in 1953.
24. W.D. Crow, personal communication.
25. F.N. Lahey and W.C. Thomas, ‘Alkaloids of the Australian Rutaceae: Acronychia baueri. I. The Isolation of the Alkaloids’, Aust. J. Sci. Res., 2A (1940), 423–426. A second part in this series was: R.D. Brown, L.J. Drummond, F.N. Lahey and W.C. Thomas, ‘II. Some Reactions of the Alkaloid Acronycine’, Aust. J. Sci. Res., 2A (1949), 622–629. Related papers were: R.D. Brown and F.N. Lahey, ‘The Ultraviolet Absorption Spectra of the Acridone Alkaloids I. Compounds Containing the Acridone Nucleus’, Aust. J. Sci. Res., 3A (1950), 593–614; Idem, ‘The Ultraviolet Absorption Spectra of the Acridone Alkaloids. II. Compounds Related to 4- Quinolone’, Aust. J. Sci. Res., 3A (1950), 615.
26. G. K. Hughes and K.G. Neill, ‘Alkaloids of the Australian Rutaceae; Evodia xanthoxyloides F Muell. I. Evoxanthine’ Aust. J. Sci. Res., 2A (1949), 429–436. G. K. Hughes, K. G. Neill and E. Ritchie, ‘The Synthesis of Melicopine and Some Trimethoxy–10- methylacridones’, Aust. J. Sci. Res., 3A (1950), 497–503. G. K. Hughes, K. G. Neill and E. Ritchie, Alkaloids of the Australian Rutaceae: Evodia xanthoxyloides F. Muell. II. Isolation of the Alkaloids from the Leaves’, Aust. J. Sci. Res.,5A (1952), 401–405. J. R. Cannon, G. K. Hughes, K. G. Neill and E. Ritchie, ‘Alkaloids of the Australian Rutaceae: Evodia xanthoxyloides F. Muell. III. The Structures of the Coloured Alkaloids, Evoxanthidine, Xanthevodine and Xanthoxoline’, Aust. J. Sci. Res., 5A (1952), 406–411.
27. CSIR became CSIRO in May 1949: C.B. Schedvin, Shaping Science and Industry: A History of Australia’s Council for Scientific and Industrial Research, 1926–1949 (Sydney, 1987), pp. 350, 355–361.
28. J.R. (Jack) Cannon in due course took up an appointment in the Chemistry Department, University of Western Australia, where he actively pursued studies in phytochemistry.
29. University of Melbourne Archives: Chemistry Staff Files 1946/196. Also op. cit. (n. 22), pp. 201, 202, 227.
30. E.R. Klein, ‘The Chemistry of Some Australian Alkaloids and Sesquiterpenes’, PhD thesis, University of Melbourne, 1951.
31. J.A. Lamberton, ‘The Chemistry of Some New Australian Alkaloids’, PhD thesis, University of Melbourne, 1950.
32. C.C.J. Culvenor, ‘Obituary: J.A. Lamberton FRACI 1925–2002’, Chem. in Aust., 69 (2002), 38–39.
33. H.F. Haynes, ‘The Alkaloids of Some Australian Plants’, MSc thesis, University of Melbourne, 1954.
34. University of Melbourne Archives: Chemistry Staff Files 1946/196; 1947/178; 1948/187; 1949/189; 1950/189; 1951/186.
35. Op. cit. (n. 12).
36. At this time the Organic Chemistry Section was part of the Chemical Research Laboratories, along with the Mineral Chemistry, Cement and Ceramics, Foundry Sands, Physical Chemistry, Chemical Physics and Chemical Engineering Sections, under the overall leadership of Dr Ian Wark.
37. Chemical Research Laboratories, Annual Report for the Year ending 30th June, 1959.
38. It is interesting to note that non-research staff were not identified in CSIRO Reports from this period.
39. Chemical Research Laboratories, Annual Report for the Year ending 30th June, 1961.
40. Chemical Research Laboratories, Annual Report for the Year ending 30th June, 1960.
41. Chemical Research Laboratories, Annual Report 1962–63.
42. Chemical Research Laboratories, Annual Report 1963–64.
43. Chemical Research Laboratories, Annual Report 1964–65.
44. C.C.J. Culvenor, personal communication.
45. John P. Shelton, personal communication.
46. Professor David Solomon, personal recollection.
47. Minister for Science.
48. The Humes business was formed in 1911 by the inventor of spun concrete pipe, Walter Hume. From the original plant in Adelaide, success of the product had led to operations being established throughout Australia and in many countries around the world. By the end of the 1920s, Walter Hume had established businesses in USA, United Kingdom, Japan, Germany, Brazil, South Africa and several Asian countries. Today, in many of these countries Hume pipe is well-known as high-quality spun-concrete pipe. In November 1988, CSR acquired Humes Ltd’s concrete products division.
49. A copy of the presentations is kept at CSIRO Molecular Science. Speakers at the Memorial Service were Dr Donald Price, Professor David Solomon, Dr Claude Culvenor, Dr Peter Wailes, Professor John Swan, Professor Roy Jackson, Mr Alan Pierce, Mr Jack Coombe, Mr Grattan Wilson, Ms Amy Tran, Ms Amina Price, Ms Janet Devlin, Ms Catherine Price, Mr Andrew Devlin and Mr Ben Price.

Bibliography

1. Hooper, P.L., Macbeth, A.K. and Price, J.R., ‘The “Hydrosulphides” of Carvone and laevo- 4-isoPropyl-Δ²-cyclohexen-1-one’, J. Chem. Soc. (1934), 1147–1150.
2. Macbeth, A.K. and Price, J.R., ‘The Action of Bases on Nitrophenylhydrazines: 2:4-Dinitrophenylhydrazine’, J. Chem. Soc. (1934), 1637–1639.
3. Macbeth, A.K. and Price, J.R., ‘The Action of Titanous Chloride on Nitrophenylhydrazones: p-Nitro- and 2:4-Dinitro-phenylhydrazones’, J. Chem. Soc. (1935), 151–153.
4. Macbeth, A.K., Price, J.R. and Winzor, F.L., ‘The Colouring Matters of Drosera whittakeri. Part I. The Absorption Spectra and Colour Reactions of Hydroxy-naphthaquinones’, J. Chem. Soc. (1935), 325–336.
5. Macbeth, A.K. and Price, J.R., ‘The Absorption Spectra of Nitrophenylhydrazines’, J. Chem. Soc. (1935), 1563–1567.
6. Macbeth, A.K. and Price, J.R., ‘The Absorption Spectra of 1:2:3-Benztriazoles’, J. Chem. Soc. (1936), 111–119.
7. Macbeth, A.K. and Price J.R., ‘The Action of Bases on Nitrophenylhydrazines. Part II’, J. Chem. Soc. (1937), 982–984.
8. Price, J.R. and Robinson, R., ‘Nitrogenous Anthocyanins. Part IV. The Colouring Matter of Bougainvillaea glabra’, J. Chem. Soc. (1937), 449–453.
9. Price, J.R. and Robinson, R., ‘A New Natural Colouring Matter of the Naphthalene Group’, Nature, 142 (1938), 147–148.
10. Price, J.R., Sturgess, V.C., Robinson, R. and Robinson, G.M., ‘Some New Anthocyanin Types’, Nature, 142 (1938), 356.
11. Price, J.R., Robinson, G.M. and Robinson R., Note: ‘Occurrence of Kaempferol in Crocus’, J. Chem. Soc. (1938), 281.
12. Lawrence, W.J.C., Price, J.R., Robinson, G.M. and Robinson, R., ‘A Survey of Anthocyanins. V’, Biochem. J., 32 (1938), 1661–1667.
13. Price, J.R. and Sturgess, V.C., ‘A Survey of Anthocyanins. VI’, Biochem. J., 32 (1938), 1658–1660.
14. Lawrence, W.J.C., Price, J.R., Robinson, G.M. and Robinson, R., ‘The Distribution of Anthocyanins in Flowers, Fruits and Leaves’, Phil. Trans. Roy. Soc. London B, 230 (1939), 149–178.
15. Price, J.R., ‘The Yellow Colouring Matter of Dahlia variabilis’, J. Chem. Soc. (1939), 1017–1018.
16. Price, J.R., Robinson, R, and (in part) Scott-Moncrieff, R. (Mrs Meares), ‘The yellow pigment of Papaver nudicaule. Part 1’, J. Chem. Soc. (1939), 1465–1468.
17. Price, J.R. and Robinson, R., ‘Dunnione, Part I’, J. Chem. Soc. (1939), 1522–1529.
18. Price, J.R. and Robinson, R., ‘Dunnione, Part II’, J. Chem. Soc. (1940), 1493–1499.
19. Lawrence, W.J.C. and Price, J.R., ‘The Genetics and Chemistry of Flower Colour Variation’, Biol. Rev., 15 (1940), 35–38.
20. Beale, G.H., Price, J.R. and Scott- Moncreiff, R., ‘The Genetics of Verbena. II: Chemistry of the Flower Colour Variations’, J. Genet., 61(1) (1940), 65–74.
21. Barber, H.N. and Price, J.R., ‘Nature of the Feulgen Reaction with Nucleic Acid’, Nature, 146 (1940), 335.
22. Beale, G.H., Price, J.R. and Sturgess, V.C., ‘A Survey of Anthocyanins. Part VII. The Natural Selection of Flower Colour’, Proc. Roy. Soc. London B., 130 (1941), 113–126.
23. Hughes, G.K., Lahey, F.N., Price, J.R. and Webb, L.J., ‘Alkaloids of the Australian Rutaceae’, Nature, 162 (1948), 233–234.
24. Price, J.R., ‘Alkaloids of the Australian Rutaceae: Melicope fareana. I. Isolation of the Constituent Alkaloids’, Aust. J. Sci. Res. A, 2(2) (1949), 249–254.
25. Crow, W.D. and Price, J.R., ‘Alkaloids of the Australian Rutaceae: Melicope fareana, II. Preliminary Examination of Melicopine, Melicopidine and Melicopicine’, Aust. J. Sci. Res. A, 2(2) (1949), 255–263.
26. Price, J.R., ‘Alkaloids of the Australian Rutaceae: Melicope fareana. IV. Some Reactions of 1-Methyl-4-quinolone-3-carboxylic Acid, A Degradation Product of the Alkaloids’, Aust. J. Sci. Res. A., 2(2) (1949), 272–281.
27. Crow, W.D. and Price, J.R., ‘Alkaloids of the Australian Rutaceae: Melicope fareana. V. Structure of the Alkaloids’, Aust. J. Sci. Res. A, 2(2) (1949), 282–306.
28. Lahey, F.N., Lamberton, J.A. and Price, J.R., ‘Alkaloids of the Australian Rutaceae. The Structure and Reactions of Acronycidine’, Aust. J. Sci. Res. A, 3(1) (1950), 155–171.
29. Price, J.R., ‘Acridine Alkaloids’, in The Alkaloids, Chemistry and Pharmacology, Manske, R.H.F. and Holmes, H.L. (eds.), Academic Press, New York, 2 (1952), 353–368.
30. McKenzie, A.W. and Price, J.R., ‘Alkaloids of the Australian Rutaceae; Glycosmis pentaphylla (Retz.) Correa’, Aust. J. Sci. Res. A, 5(3) (1952), 579–580.
31. Haynes, H.F., Nelson, E.R. and Price, J.R., ‘Alkaloids of the Australian Rutaceae; Pentaceras australis Hook F.I. Isolation of the Alkaloids and Identification of Canthin-6- one’, Aust. J. Sci. Res. A, 5(2) (1952), 387–400.
32. Haynes, H.F., Nelson, E.R. and Price, J.R. (1952) Alkaloids of the Australian Rutaceae: Pentaceras australis Hook F. II. Identification of 5-Methoxycanthinone’, Aust. J. Sci. Res. A, 5(3) (1952), 563–569.
33. Nelson, E.R. and Price, J.R., ‘Alkaloids of the Australian Rutaceae: Pentaceras australis Hook F. III. Identification of 4-Methylthiocanthin-6-one’, Aust. J. Sci. Res. A, 5(4) (1952), 768–781.
34. Cannon, J.R., Hughes, G.K., Price, J.R. and Ritchie, E., ‘The Chemical Constituents of Australian Flindersia Species. IV. The Constituents of Flindersia bourjotiana F. Muell.’, Aust. J. Sci. Res. A, 5(2) (1952), 420–422.
35. McKenzie, A.W. and Price, J.R., ‘The Alkaloids of Gyrocarpus americanus Jacq.’, Aust. J. Chem., 6(2) (1953), 180–185.
36. Lamberton, J.A. and Price, J.R. ‘Alkaloids of the Australian Rutaceae: Acronychia baueri Schott. IV. Alkaloids Present in the Leaves’, Aust. J. Chem., 6(1) (1953), 66–77.
37. Lamberton, J.A. and Price, J.R., ‘Alkaloids of the Australian Rutaceae: Medicosma cunninghamii Hook F.’, Aust. J. Chem., 6(2) (1953), 173–179.
38. Price, J.R., ‘Recent Developments in the Study of the Chemistry of Australian Plant Products’ (Liversidge Lecture), Rept. Australian and New Zealand Assoc. Advancement Sci. (29th Meeting, Sydney, 1952), 29 (1953), 67–80.
39. Johnstone, R. and Price, J.R., ‘N-Chloroacetylisatic Acid’, Aust. J. Chem., 7(2) (1954), 209–210.
40. Culvenor, C.C.J., Drummond, L.J. and Price, J.R., ‘Alkaloids of Heliotropium europaeum L. I. Heliotrine and Lasiocarpine’, Aust. J. Chem., 7(3) (1954), 277–286.
41. Price, J.R., ‘Plant Chemistry in Australia’, Rec. Chem. Progr., 16(3) (1955), 153–163.
42. Price, J.R., ‘Structure of Lunamine’ (title only, paper read at Section B), Rept. Australian and New Zealand Assoc. Advancement Sci. (31st Meeting, Melbourne, 1955), 31(1955), 47.
43. Price, J.R., ‘Alkaloids Related to Anthranilic Acid’, Fortschr. Chem. Org. Naturst., 13 (1956), 302–345.
44. Price, J.R. and Smith, L.W., ‘The Reaction of N-Chloroacetylisatin with Alkali’, Aust. J. Chem., 9(1) (1956), 139–140.
45. Price, J.R., ‘Some Aspects of Organic Chemical Research as Applied to Agriculture’, J. Aust. Inst. Agr. Sci., 22(1) (1956), 3–10.
46. Price, J.R., ‘A Novel Type of Naturally Occurring Quaternary Base’. In Current Trends in Heterocyclic Chemistry, Proc. Symp. Canberra, 1957 (Albert, A., Badger, G.M. and Shoppee, C.W., eds.), Butterworths, London (1958), 92–109.
47. Johnstone, R., Price, J.R. and Todd, A.R., ‘Alkaloids of the Australian Rutaceae; Lunasia quercifolia. I. 7-Methoxy-1-methyl- 2-phenyl-4-quinolone’, Aust. J. Chem., 11(4) (1958), 562–574.
48. Price, J.R., ‘Alkaloids of the Australian Rutaceae: Lunasia quercifolia. II. The Nature of Lunasine’, Aust. J. Chem., 12(3) (1959), 458–467.
49. Price, J.R. and Willis, J.B., ‘The Infra-red Spectra of 2- and 4-Quinolones’, Aust. J. Chem., 12(4) (1959), 589–600.
50. Price, J.R., ‘Australian Natural Product Research’, Pure Appl. Chem., 2 (1961), 367–381.
51. Baldwin, M.E., Bick, I.R.C., Komzak, A.A. and Price, J.R., ‘Some Ketones from Acradenia franklinii’, Tetrahedron, 16 (1961), 206–211.
52. Price, J.R., ‘The Distribution of Alkaloids in the Rutaceae’. In Chemical Plant Taxonomy (Swain, T., Ed.), Academic Press, London, 1963, 429–452.
53. Price, J.R., ‘Home Thoughts from Abroad’. Proc. Roy. Aust. Chem. Inst., 30 (1963), 89–93.
54. Price, J.R., ‘The Future of the Institute’, Proc. Roy. Aust. Chem. Inst., 30(8) (1963), 297–309.
55. Price, J.R., ‘Studies in Alkaloids’, Farmaceutisk Revy, 62 (1963), 145–155.
56. Price, J.R., ‘Antifertility Agents of Plant Origin’. In Agents Affecting Fertility, Symp. London, 1964 (Austin, C.R. and Perry, J.S., eds.), Churchill, London, 1965, 3–16.
57. Hart, N.K. and Price, J.R., ‘Alkaloids of the Australian Rutaceae: Lunasia quercifolia. III. Isolation of (-)-O-Methylluninium Salts’, Aust. J. Chem., 19(11) (1966), 2185–2187.
58. Inubushi, Y., Sano, T. and Price, J.R., ‘Triterpene Constituents of Lycopodium complanatum L. from New Guinea’, Aust. J. Chem., 20(2) (1967), 387–388.
59. Lamberton, J.A., Price, J.R. and Redcliffe, A.H., ‘Micromelin, a New Coumarin from Micromelum minatum (Forst.f.) Seem (Family Rutaceae)’, Aust. J. Chem., 20(5) (1967), 973–979.
60. Johns, S.R., Lamberton, J.A. and Price, J.R., ‘(±)-N-Benzoyl[2-hydroxy-2(4’-methoxyphenyl)] ethylamine from Clausena brevistyla Oliver (Family Rutaceae)’, Aust. J. Chem., 20(12) (1967), 2795–2797.
61. Price, J.R., ‘The Organisation of Chemists and Chemistry’, Proc. Roy. Aust. Chem. Inst., 34(9) (1967), 239–241.
62. Hart, N.K., Johns, S.R., Lamberton, J.A. and Price, J.R., ‘Alkaloids of the Australian Rutaceae: Lunasia quercifolia. IV. Identification of a Minor Constituent as 5-Hydroxy-1- methyl-2-phenyl-4-quinolone and Preparation of an Angular Isomer of (-) Lunine’, Aust. J. Chem., 21(5) (1968), 1389–1391.
63. Johns, S.R., Lamberton, J.A., Price, J.R. and Siomis, A.A., ‘Identification of Coumarins Isolated from Lepiniopsis ternatensis (Apocynaceae), Pterocaulon sphacelatum (Compositae) and Melicope melanophloia (Rutaceae)’, Aust. J. Chem., 21(12) (1968), 3079–3080.
64. Johns, S.R., Lamberton, J.A. and Price, J.R., ‘Isolation of Isomultiflorenol, a Possible Triterpenoid Artefact, from Pleiococca wilcoxiana (Rutaceae)’, Aust. J. Chem., 23(6) (1970), 1283–1284.
65. Price, J.R., ‘The Communication of Scientific Knowledge for Useful Application’. The 1970 Leighton Address. Proc. Roy. Aust. Chem. Inst., 38(5) (1971), 113–121.
66. Price, J.R., Graduation address. Australian National University, Canberra, April 1971. Australian National University News, 6:(2) (1971), 15.
67. Price, J.R., New Inventions and Riches. Australian Director, 2:(2) (1972), 13–16.
68. Price, J.R., Report on 28th Annual Conference. Opening remarks. Appita Journal, 27:(6) (1974), 382–383.
69. Barnes, C.S., Price, J.R. and Hughes, R.L., ‘An Examination of Some Reputed Antifertility Plants’, Lloydia, 38 (1975), 135–140.
70. Price, J.R., CSIRO: Fifty Years of Research. Looking to the Future. Nature, 261:(5562) (1976), 631–632.
71. Collins, D.J., Culvenor, C.C.J., Lamberton, J.A., Loder, J.W. and Price, J.R., ‘Plants for Medicines: A Chemical and Pharmacological Survey of Plants in the Australian Region’, CSIRO Publishing, Melbourne, 1990.
72. Price, J.R., Lamberton, J.A. and Culvenor, C.C.J., ‘The Australian Phytochemical Survey: Historical Aspects of the CSIRO Search for New Drugs in Australian Plants’, Historical Records of Australian Science, 9(4) (1993), 335–356.

Written by Ian Franklin, Geoff Grigg and Oliver Mayo.

• Life
• Work
• Leadership
• Later years
• About this memoir
  • Acknowledgments
  • References
  • Publications of J.M. Rendel
  • Notes

Life

James Meadows Rendel was born on 16 May 1915 in England. He moved to Australia in 1951 to join CSIRO and was appointed Chief of the Division of Animal Genetics in 1959. He was elected to the Australian Academy of Science in 1960, retired from CSIRO in 1980 and died on 4 February 2001. His influence on genetics and the development of the theory and practice of animal breeding in Australia was profound.

After his election to the Academy, Rendel served on its Council in 1963-5 and 1971-4, being a vice-president in 1973-4. He was Burnet Lecturer in 1981.

Rendel's family background (important to understanding an Englishman) was intellectual. Although his father, Colonel Richard Meadows Rendel, was a professional soldier, there were connections to the Bloomsbury group on both sides: his father's uncle was Lytton Strachey, and his mother's sister was the diarist Frances Partridge. Rendel's first marriage, on 22 October 1938, was to the poet Joan Adeney Easdale, a protégée of Leonard and Virginia Woolf who published some of her poems in their Hogarth Press. Friends included Professor Lionel Penrose the human geneticist (whose brother was a champion of cubism and cubists in England) and the Mitchison clan. As a student, Rendel played the flute, was fascinated by ballet and kept a chameleon. He received part of the normal education of an upper- middle-class Englishman of his time (Rugby School), but went to University College London rather than Oxbridge and completed his PhD as a student of J. B. S. Haldane, the great geneticist and left-wing science popularizer. Working with Haldane gave him a breadth as a scientist that was a great boon throughout his career, but also left him in due course with a permanent physical disability. We believe that Rendel and John Maynard Smith, another notable evolutionary geneticist, were Haldane's only PhD students, something that seems unlikely and which we have been unable to confirm.

During the Second World War, when Rendel was attached to RAF Coastal Command, he took part in Haldane's experiments on escape from submarines (critical after a peace-time disaster). Haldane never spared himself as a guinea-pig, and those of his colleagues with as much courage took the same risks. One experiment involving a high compression chamber was disastrous and almost fatal for Rendel, as both of his lungs were punctured and he was left with permanent pulmonary problems. General details of this work can be found in Ronald Clark's biography of Haldane (Clark 1968), but it is characteristic of Rendel, who helped Clark considerably with the book, that the reference to this accident does not mention Rendel by name. As most of the actors in these events are no longer alive, Clark's book remains the best published source, incomplete though it is.

After the war, Rendel moved to Edinburgh to join the legendary animal genetics research group put together in the University of Edinburgh by C. H. Waddington, a man of such huge ego that he never hesitated to appoint better scientists than himself to his Institute. Two such were Jim Rendel and Alan Robertson, both of whom were known to Waddington from his time as Chief of Operational Research in Coastal Command. (They had worked on methods for detecting German U-boats at sea.) They were put in charge of a dairy research programme, and together they made several vital advances in the design and analysis of dairy breeding programmes, establishing principles that laid the foundation for the widespread use of artificial insemination in dairy progeny testing programmes. Later, Rendel implemented these ideas in CSIRO's tropical dairy breeding initiative. Later still, Robertson (1977) wrote a biographical memoir of Waddington that stresses canalisation and yet, perhaps at first sight surprisingly, does not mention Rendel. However, Robertson did not agree with Rendel's approach to Drosophila bristle epigenetics (of which much more below), and to mention his old friend would have been to criticize him.

Rendel was then recruited from Edinburgh in 1951 by Sir Ian Clunies Ross, Chairman of CSIRO, to set up a CSIRO group at the University of Sydney to teach animal genetics and to develop and supervise a programme of research into animal breeding methods encompassing the domestic fowl, sheep, dairy and beef cattle. Rendel's recruitment, along with those of Otto Frankel (CSIRO, Canberra) and David Catcheside (University of Adelaide), was the culmination of an intensive effort by Clunies Ross to re-establish the science of genetics in Australia following its decline in the pre- and post-war years (see McCann and Batterham 1993)

When he arrived from Edinburgh, Rendel and the two colleagues he brought with him were housed in premises at the University of Sydney where they began to teach genetics in both the Science and Veterinary Science Faculties. As he developed his Section, it grew in size and responsibilities and CSIRO made it an independent Division, of Animal Genetics, in 1959.

His son Sandy Rendel notes that 'Jim really liked the farmers he came in contact with. On the boat out (the P & O liner Oronsay) he and my mother became friends with Charles and Amy Cooper. Charlie Cooper owned Kunanadgee, 3000 acres on the River Murray at Corowa. It was one of the farms Jim visited on his familiarisation tour after arriving in Australia. Subsequently it was involved in the release of myxo.[1] I spent a lot of my school holidays there. Jim also enjoyed his trips to the research station at Cunnamulla and working with Bill Gunn and the meat board' (S. Rendel, pers. comm., 2004)

Whilst he enjoyed his involvement in animal breeding – and cattle breeding was a passion – Rendel's primary interest was in how genes worked in the animal. In his first year in Australia, he joined with a number of other geneticists to found the Genetics Society of Australia. He set up groups in the Section of Animal Genetics to study the fundamentals of genetics, using mice, Drosophila and Paramecium as experimental organisms. In the 1960s, he realised the future impact that molecular genetics must have in animal improvement, and set out to establish molecular biology in the Division. This group later was spun off as CSIRO's Molecular and Cellular Biology Unit. He had a particular personal interest in the genetics of developmental processes in the whole animal (here the influence of Waddington during his five years in Edinburgh is evident). Rendel explored and wrote about novel ideas on developmental 'canalisation', discussed in detail in the next section; his only published book, Canalisation and Gene Control, covers the topic.

Although Rendel continued his own research with Drosophila, and built new research programmes in molecular and developmental biology within his own Division, he recognised his responsibility to provide 'something for now, something for later' (in Justus von Liebig's words) very broadly, and he had the vision of, and the resources to realise that vision in, new breeds as well as new scientific concepts.

In the application of genetics to animal breeding, Rendel's main personal contribution was to the development of new breeding programmes, especially for northern Australia. CSIRO had already imported Indian and African cattle to develop a livestock industry for tropical and sub- tropical regions, and Rendel encouraged basic research into understanding the interactions between adaptation to stressful environments and productivity in those environments. The improvement of adaptation in beef cattle by use of African germplasm resulted in the Belmont Red; in dairy cattle, Indian cattle and European dairy cattle were used to produce the Australian Milking Zebu (AMZ). Had typical Australian parochialism not led to a rival programme in Queensland that produced a competing breed, the Australian Friesian Sahiwal (AFS), Australia might have led the tropical world in developing a sustainable dairy breed and production system. In contrast, the beef cattle industry in Australia's north has been a major success. In poultry, Rendel's ideas on the practical application of canalisation theory to decrease the inter-egg interval in layers made, through Bruce Sheldon, a major contribution to the egg industry before that industry closed genetics research in favour of importing germplasm.

A new tropical beef cattle research laboratory was built in Rockhampton at about the time Rendel retired as Chief of Division, and it was named the James Rendel Laboratory to recognise his contribution. There is no comparable memorial to his contribution to sheep breeding, where, with his quiet support and critical direction, Helen Newton Turner and others pioneered objective measurement of fleece traits and struggled to help sheep breeders, in many cases against their will, to apply the same successful science to the wool industry as had revolutionized pigs, poultry and dairy.

He knew the importance of encouraging young scientists and was always generous with his time and CSIRO's resources to help those whom he judged deserving. Stuart Barker, Jim Peacock and the authors of this memoir are among the many who benefited from his quiet early encouragement. Under his leadership, the Animal Genetics section was egalitarian and a fount of ideas that reverberated through the emerging Genetics community. His arguments were always stimulating but often obscure, especially to those who saw animal breeding as a science rooted in statistics rather than biology.

Upon his retirement from CSIRO in 1980, Rendel briefly took up a visiting professorship at Harvard to collaborate with his old friend Richard Lewontin. Subsequently he and his wife Tresham moved to live in a fine house on a smallholding at Drinkstone Green in Suffolk, England, where he bred Booroola sheep which he had imported from Australia. This was following in his father's footsteps, for Colonel Rendel had retired early to raise poultry in Kent, returning to active service for the Second World War.

Australia, however, continued to beckon and seven years later he and Tresham returned to Sydney and to Wentworth Falls in the Blue Mountains above Sydney where they settled. An old friend and colleague, Bill Sobey, writes that in his opinion Rendel did not 'ever become an Australian', he was too English, but he had a deep affection for his adopted land.

In this second retirement, Rendel maintained his lifelong interest in thinking and writing about science, and wrote an entertaining and challenging book about the role of common sense in judging science.

He was survived by his second wife Tresham (Marie Tresham Davies; Tresham was a family name, one of her ancestors being Francis Tresham, a co-conspirator with Guy Fawkes in the English Gunpowder Plot of 1605), his six children (Jane Susan Robertson, Polly Mary Virginia Woods, Alexander Meadows Rendel, twins Julia Margaret Szulerowski and Richard James Rendel and Francis Kate Hayes), ten grandchildren and one great-grandchild.

Work

Early work and influences

Rendel's earliest work, as a PhD student in London, was a study of the relationship between egg size and hatchability in ducks. He published a number of papers arising from this work, demonstrating in particular that intermediate egg weights are favoured by natural selection. Undoubtedly, this work drew his attention to mechanisms that allow for reduced variability, a topic he revisited later in his Drosophila experiments. Later, when he joined Waddington's group in Edinburgh, he was drawn to the application of genetics to animal breeding and formed a close association with Alan Robertson. Together, Rendel and Robertson formulated a set of protocols for dairy improvement based on progeny testing and the widespread use of artificial insemination. This work was seminal in the design and analysis of dairy breeding programmes, especially in Europe, and their methodologies were in many ways superior to those being developed at the same time by Hazel and Lush in the USA. Their work led to a decline in the use of testing stations and the widespread use of family selection in selecting dairy bulls. When Rendel arrived in Australia, plans to develop tropically adapted dairy cattle were already under way through the efforts of R. B. Kelley, and Rendel grasped the opportunity to apply the principles developed by him and Alan Robertson to design the first animal improvement programme in Australia based on modern quantitative genetics theory. While the AMZ programme, as it later became known, has not been a commercial success, the principles of well designed progeny testing programmes had a very considerable influence on early dairy improvement programmes in Australia. In parallel, Rendel did much to set the direction of another animal breeding programme for the tropics, based at Rockhampton and directed at the beef industry.

However, perhaps the greatest influences on Rendel while in Edinburgh were the ideas of Waddington. It was here that Rendel developed a deep commitment to the importance of developmental biology and in particular Waddington's ideas on canalisation and genetic assimilation that led to Rendel's work on the canalisation of bristles in Drosophila. Finally, Rendel formed a strong relationship with Alex Fraser. Fraser, a New Zealander and a friend of Otto Frankel, had arrived unannounced in Edinburgh to do a PhD with Waddington. Rendel encouraged Fraser to join him in Sydney when Fraser had completed his PhD. Fraser arrived slightly before Rendel and began to set up the animal genetics unit in the Division of Animal Health and Production, in preparation for Rendel's arrival. Fraser's work on wool biology, and later in Drosophila, influenced Rendel to consider interactions among the components or determinants of any production trait, such as fleece weight. One of Rendel's later important publications, with Ted Nay, reflects those influences.

Major achievements

Much insight into Rendel's scientific outlook and approach, as well as knowledge of his results, can be obtained from Canalisation and Gene Development, which is based on a series of lectures and is in consequence clear and direct. Rendel stakes his claim in the first sentence of the preface: 'This book treats development as though it were a process initiated by a major gene and regulated through the major gene's action.' (p. 9). The experimental approach involved was not novel, in that many others (e.g. Grüneberg in mice) had studied the disruptive effects of abnormal alleles of major genes in order to understand development, following the great William Harvey's advice that

Nature is nowhere accustomed more openly to display her secret mysteries than in cases where she shows traces of her workings apart from the beaten path; nor is there any better way to advance the proper practice of medicine than to give our minds to the discovery of the usual law of Nature by careful investigation of rarer forms of disease. For it has been found in almost all things, that what they contain of useful or applicable nature is hardly perceived unless we are deprived of them, or they become deranged in some way.

We have included this lengthy quotation from Harvey (1657; 1847) because it expresses so much of Rendel's own attitude. Rendel's work was informed by the philosophy set out by Waddington (1957) but instilled into his colleagues in Edinburgh over many years.

At the time he began his work, Rendel could not hope to isolate and describe the genes involved in the control mechanisms of development, so he placed his emphasis on describing and measuring how a particular major gene influenced development of a particular set of traits. He sought to describe aspects of the process as a functional relationship between some underlying gene outcome or product, M, and the trait or phenotype, P, that is, P = f(M). He called this underlying 'complex of influences taking part in the making of a phenotype', Make (M). It is not a term that has taken on, in contrast to Waddington's canalisation, the regulation of development that produces a normal outcome in the face of environmental shocks, which is widely recognised today as important with well developed theory and experimental support, including Rendel's own pioneering work (see e.g. Gibson and Wagner 2000 and Kitami and Nadeau 2002)

It is possible that Rendel was unfortunate in the timing of his fundamental as opposed to his applied work, since it is not based on attack on the problem of the nature of the gene – that is, on how DNA constitutes the genetical message, which was the fashion throughout his civilian working life – whereas developmental genetics came into its own only during his final decade. He was certainly unfortunate in trying to quantify development, through investigation of f(M), in an era when numeracy was not demanded of developmental biologists. Many found probits (Fisher and Yates 1963) rebarbative.

Rendel investigated a sex-linked gene, scute (sc), that influenced the formation of bristles on the scutellum of Drosophila melanogaster. In the wild type, there are normally four bristles in this location, and the variance about this number is very low; in Waddington's term, scutellar bristle number is canalised at four. The sc gene both reduces the number of bristles formed and increases the variance of this number. Selection for increased bristle number in animals carrying one or more sc genes produced an increased number of bristles in wild-type flies as well as those carrying one or more sc.

On the assumption that underlying variation in M is Gaussian in distribution, the probit transformation allows estimation of intervals of M corresponding to frequency classes of different numbers of bristles (0,1,2,3,4,5,...). In this way, Rendel could show that a large range of M yielded four bristles in wild-type flies, thereby quantifying the canalisation, whereas outside this range, small changes in M could produce large changes in P. Thus, the shape of f(M) could be and was determined experimentally.

A second important finding, which confirmed Fisher's work on the evolution of dominance (though Rendel disagreed with Fisher in his interpretation of Fisher's experiments; cf. Fisher 1999), was that selection of modifiers of bristle number was possible through the disruptive effect of the major gene, but the selected modifiers were not regulated by the same system, or they would not have been selectable. Rendel concluded that dominance, though a primary concept by virtue of its recognition by Mendel a century previously, was not primary in a biological sense: developmental stability was the outcome of natural selection, and dominance of the wild type a threshold effect that was a consequence of canalisation.

Rendel, having explained evolved dominance and other phenomena in this fashion, went on to consider the action of selection more generally. His discussion of the different types and consequences of selection for an intermediate phenotype is a model of clarity. One at least of his conclusions remains important: regulation is an outcome of natural selection on account of its benefits to developmental processes, not because it yields intermediate optima. From his own experimental work and a fair-minded evaluation of that of others, he showed that variability about an intermediate value could be successfully reduced by selection if that selection was carried out among animals whose variability was enhanced by a major gene such as sc, but not otherwise. He noted in contrast that many traits were not canalised, in that directional selection, whether for an increase or a decrease, was always successful. In consequence, the variability that is important in plant and animal breeding should always be assessed for canalisation before selection was undertaken.

Rendel attempted to relate his quantitative schema to gene regulation as it was understood at the time, in particular to Jacob-Monod operon theory, but the two approaches were too far apart to yield valuable results. He was closely in touch with the development of metabolic control theory by Henrik Kacser and others, and pointed out that the case of sc could not be fitted into Kacser's framework. Recently, Wagner and colleagues (Bagheri-Chaichian and Wagner 2002, Bagheri-Chaichian, Hermisson, Vaisnys and Wagner 2002, Wagner, Booth and Bagheri-Chaichian 1997) and Omholt, Plahte, Øyehaug and Xiang (2000) have brought this discussion up to date and have shown that homoeostasis and hence canalisation are not inevitable. That is, phenomena like canalisation and dominance may be the outcome of evolution by natural selection. In the case of dominance, it may arise as an ancillary outcome of direct selection on traits controlled by genes that are likely to influence dominance. In the case of canalisation, there is an interaction between direct stabilizing selection on a trait and selection for canalisation of that trait, such that if the genetic variance in the trait is reduced to a very low level, canalising selection will be ineffective. Rendel's contribution, which pointed qualitatively towards many of these conclusions, has been largely absorbed in time.

The final stage of Rendel's work was its practical application, the Eggatron being the most important example.[2]

This was a daylight-excluding poultry layer-house with automatic recording of the time of lay for every hen and the capability to vary day length as experienced by the hens. In nature, hens lay eggs daily for a number of days, yielding a clutch, and then set this clutch to hatch. Before Rendel's work, increased egg numbers had come first from the breaking of the link between laying and setting, then from reduction of the inter-clutch interval and then from increase in the number of consecutive clutches, though these stages were not necessarily recognized as separate or separately selected. There may also have been selection for clutch size, but this is often highly canalised (see e.g. Mayo 1983, Chapter 7), and we know little of clutch size in ancestral poultry. In the Eggatron, day length less than 24 hours exposed additional genetical variation in rate of lay, so that the interval between eggs could be reduced from 24 hours. Increased rate of lay was obtained by Rendel's colleagues, working to his plans. At one time, 40% of Australian layer germplasm came from the Eggatron, through dedicated application of Rendel's ideas by Bruce Sheldon and others. Sadly, commercial breeders later chose to import germplasm rather than continue advanced work based on CSIRO research, condemning the Australian industry to import replacement and export uncompetitiveness, but Rendel's approach was commercially successful.

Helen Newton Turner's work on the Booroola gene for increased fecundity in sheep was a parallel development that was as scientifically sound and yielded, through its application to meat sheep by L. R. A. (Laurie) Piper and B. M. (Bernie) Bindon, potential for increased lambing, but it has not had as yet the commercial success of the Eggatron.

Leadership

We have already mentioned some of the people who influence Rendel in his research directions, in particular Haldane and Waddington. Haldane and his family upbringing made Rendel respect intellect deeply but be no respecter of persons – authorities in particular. Though autocratic in some ways, he yet insisted that scientists must have intellectual freedom and adequate resources to pursue their ideas.

As we have already noted, Waddington had enormous, indeed unlimited, regard for his own abilities and was delighted to appoint staff who were, as it turned out, better scientists than he, such as Alan Robertson. A man of very broad interests to whom art and travel and the company of congenial intellectuals were hugely important, he never had time to supervise his staff closely, and by appointing those whom he knew and trusted from his wartime work, as well as brilliant young people in related fields, he created a scientific powerhouse in the Institute of Animal Genetics. From him, Rendel learned to appoint the most talented people he could find, to give them very general direction, and to let them work towards success or failure. The results, as might have been predicted, were mixed.

Rendel was appointed to head the new Division of Animal Genetics in a golden hour for genetics, for CSIRO, and for the industries that the Division was to serve. His success or otherwise was therefore dependent on his overall vision, and his ability to choose the people who would bring it to fruition.

CSIRO had been established in 1949 from the already successful CSIR and was widely respected for its achievements in the national interest. Indeed, in the public mind it was almost synonymous with science, and its activities were the major part of Australia's non-defence research from 1926 on (see Mayo 2002 for discussion and references). Its founding Chairman, Ian Clunies Ross, had access to everyone from the Prime Minister down, and though he died the year before Rendel took up his post as a Divisional Chief, CSIRO's standing, and consequently its funding, were unquestioned for many years.

Rendel's Division was strongly supported by the wool and beef industries, though not as strongly as was the Animal Physiology Division by wool. The history of the wool industry in the last half-century makes gloomy reading, and many have argued that the failure of breeders to take up methods proven successful in the pig and poultry industries was in part a failure of the applied-science leadership to take the work out to industry wholeheartedly and coherently. Rendel never saw technology transfer as one of his Division's primary responsibilities, so that more effort was put into this activity in tropical dairy breeding because otherwise no progress could have been achieved, than in wool breeding. State Departments of Agriculture and Primary Industry unquestionably had technology transfer, or extension as it was usually called, as part of their mandate, and Rendel approved of and supported strong collaboration between his Division and these agencies, but he never led the effort himself. He saw his role as scientific leadership, and in this role he was fearless.

While animal production research was to some extent established in the Division of Animal Health and Production by the time that Jim Rendel arrived, he had built upon it steadily in the 1950s. Arthur Dunlop had come to the Division from a PhD at Iowa State University to carry out wool research, and Helen Newton Turner was appointed to lead sheep genetics research in the Division. H. G. Turner was recruited to head a small beef research group in Rockhampton to study the relationship between production and adaptation to tropical environments. A property, Belmont, was purchased with support from the Australian Meat and Livestock Corporation to provide experimental material for this research. Later, a poultry research group, based initially at Werribee, was moved to Sydney in new facilities at North Ryde. At Badgery's Creek, south-west of Sydney, two zebu milking breeds, the Sindhi and Sahiwal imported from Pakistan, were crossed to Jerseys under the supervision of Bob Hayman. Geoff Grigg and Hymie Hoffman were moved to Sydney from Adelaide to establish molecular and developmental biology research. Alex Reisner, who had worked on Paramecium, joined the Division, as did Peter Claringbold, who subsequently rose to the position of Chief of Computing Research.

In 1959 the Division of Animal Health & Production was split into three Divisions, Animal Health, Animal Genetics (under Rendel), and Animal Physiology at Prospect (a site initially set up by Harold Carter as a wool research laboratory). By the late 1960s, Rendel had recruited additional molecular biologists, notably Hiro Sibatani and Stephen Fazekas de St Groth, as he had developed a firm conviction that the future of genetics research lay in molecular and developmental biology.

Rendel's leadership style was based on the principles of the Duke of Wellington – who believed in appointing the best people that he could find, and then not interfering with their decisions – as put into practice by Waddington. Martin (1997, Chapter 2) gives a view of one case where this did not work, but within the Division it was generally highly successful. As CSIRO grew, its procedures became more formal (some would say bureaucratic), especially in the early seventies, and Rendel had difficulties with the senior management of CSIRO because he refused to tolerate interference or calls to justify his decisions. The Division was disbanded in 1975, with the more traditional areas merged with the Division of Animal Physiology and the molecular and genetics research spun off as a separate unit, later the Division of Molecular Biology. This was unfortunate, and was based at the time on a report by Eric Underwood, who saw a need for geneticists and physiologists to spend more time talking to one another, but who saw no relevance of molecular biology to animal production research. Rendel remained with the newly formed Division of Animal Production, returning to the bench until his retirement in 1980. His outstanding scientific leadership capabilities, however, were no longer used by CSIRO.

Initially, Rendel's laboratories were centred on the University of Sydney – a model of collocation that still works very well – with field stations both nearby at Badgery's Creek, at that time only an hour's drive west of Sydney, and far away at Gilruth Plains near Cunnamulla in south-western Queensland. The mandate was the national benefit. In that great era when Australia recognised the need to expand its research effort, resources were made available to bring the basic science and its applications to sheep and poultry together, and new premises were opened on the CSIRO North Ryde campus in 1963. The poultry genetics unit was moved to North Ryde from Werribee in 1965-66. The new Division now included branch laboratories and field stations at Rockhampton (tropical beef cattle), Armidale (sheep), and Wollongbar (tropical dairy cattle) as well as those already mentioned.

The appointments Rendel made were diverse and remarkable. Some have already been discussed. We do not discuss them all, and we exclude ourselves.

Two of the earliest appointments were Alex Fraser and Bill Sobey, both colleagues in Edinburgh. Alex Fraser (elected FAA but later resigned on relocating permanently to the USA) conducted many valuable fundamental experiments on pattern formation in such cases as bristle number in mice. He contributed powerfully to the discussion of any and all topics at seminar and in the tea room. His monograph with Short on the biology of the fleece was an important piece of work, but it was not taken up by the 'practical' geneticists led by Helen Newton Turner. The failure of most of the developmental biologists to engage directly in Merino fleece genetics and breeding was a background reason for the divergence in approach in the Division between those who wanted to apply basic quantitative genetics to fleece weight and fibre diameter to 'fine the national clip' and those who wanted a subtler but inevitably slower approach that required the interactions among the components of fleece weight to be experimentally and theoretically elucidated before fining the clip.

Bill Sobey, a personal friend and colleague, carried out significant work on rabbit fleas as vectors for myxomatosis. In 1975, following the merger of Animal Genetics with Animal Physiology, he transferred to Wildlife Research to continue important work that was no longer in the production sphere.

Peter Claringbold was a veterinarian turned computer scientist, not a particularly unusual transition at a time when no computer scientists per se were being trained. Before becoming Chief of CSIRO's newly established Division of Computing Research, he was one of many who contributed to the pioneering computerization of both breeding programmes and general record-keeping for the poultry and dairy breeding programmes. He assisted Alex Fraser enormously in his early work on computer simulation of genetic systems.

H. G. Turner, never to be confused with Helen Newton Turner, led the cattle- breeding work at Rockhampton after Rendel. He was a very clear thinker, did much important work on the nexus between adaptation and productivity in the tropics, and led the development of an internationally recognised tropical cattle research centre. Turner was ably followed by John Vercoe. Both are examples of capable scientists who did not consider that good work had to be done in a metropolitan centre; without their kind, the cattle industry in northern Australia could not have become the success it is today, nor could there have been continuity in the more basic work on mechanisms of heat tolerance, tick resistance and other important traits. The Australian Academy of Technological Sciences and Engineering has identified CSIRO's tropical beef breeding as one of the major successes in agricultural technology in Australia's first 200 years of European settlement (AATSE 1988).

Emeric Binet, part of the Hungarian diaspora that has so much enriched Australia's social, intellectual and business life since the Second World War, was a mathematician who had turned his hand to genetics. Unfortunately, brain damage suffered in a motor accident made it hard for him to concentrate on work, though the force of his intellect was undiminished. Characteristically, Rendel did not seek invalidity for Binet, believing that he could still contribute through discussion. This happened, but Binet's major contribution for many was as a source of legends, such as those to do with the consequences of damage to his thermoregulatory centre. Emeric was but one of many mathematical geneticists appointed to the Division. Pre-eminent, of course, was Helen Newton Turner, whose role in the development of sheep-breeding research was pivotal and is well known in the industry. Helen very much ran her own group within Animal Genetics, ably assisted by many in her group. The most important of her colleagues, perhaps, were Arthur Dunlop and Sid Young.

Complementing the growing strength of the Division in developing animal-breeding methodologies, Rendel's efforts to strengthen molecular and developmental biology, both in his Division and in Australia, resulted in the appointments of Grigg, Hoffman and Reisner, each of whom worked in various aspects of molecular and cellular biology. Additional appointments were made in the mid-sixties.

Hiro Sibatani joined Animal Genetics as a molecular biologist. Japanese, he was an extraordinarily talented yet slightly eccentric figure whom most of his colleagues loved. He was even more interested in the Philosophy of Science than Binet and his philosophical contributions to laboratory and tea-time discussion were more profound and productive than Binet's.

Stephen Fazekas de St Groth (FAA) was a Hungarian molecular biologist recruited specifically by Rendel to bolster molecular research in the mid-sixties. He was outstandingly able and conducted important work on the influenza virus, but did not have a major impact on animal production science. He helped establish a number of techniques in the Division, such as the development and application of monoclonal antibody technology.

Many others have made important contributions to Australian science. Among these are Bruce Sheldon and Brian Yoo, who led the poultry genetics programme, Bernie Bindon and Laurie Piper, who worked together on the genetics of fecundity in sheep, and later came to lead Co-operative Research Centres in beef cattle and sheep research, and Judith Koch, a molecular biologist working closely with Stephen Fazekas.

As we have already indicated, the atmosphere at North Ryde was intellectually exciting. Morning tea could turn into a seminar, seminars could turn into non- violent pitched battles of argument, everyone was open to challenge about his or her work at any time, except perhaps during the chess games that were a permanent feature of the tea room. Rendel himself was a keen chess and (contract) bridge player, or indeed participant in anything that sharpened one's wits.

Later years

Rendel retired from CSIRO in 1980, as already mentioned. At that time retirement at 65 was mandatory, regardless of the officer's capabilities and inclinations. Rendel's intellectual vigour and enthusiasm were as great as ever, and he was in demand as a consultant and adviser on cattle breeding, as his publication list shows.

He had always been a wide reader: his children remember him reciting 'The love song of J. Alfred Prufrock' from memory, and his interests ranged from Wellington (his wife Tresham gave him a complete set of Wellington's despatches, which he greatly enjoyed), through the King James Bible to Gibbon, Wells, Virginia Woolf and Russell. He was particularly interested in the grand sweep of history of men and ideas.

He was fascinated by the problems of consciousness: its nature and its relationship to the material world. He was hostile to the idea that new properties emerged as the complexity of organization increased in the physical world ('emergentism') and was therefore led to the belief that consciousness, albeit in a primitive form, must be as fundamental to the universe as gravity or mass. This view is philosophically unfashionable but in the view of Rendel's son-in-law Sandy Robertson has interesting parallels in the work of the Australian philosopher David Chalmers (e.g. Chalmers 1996).

His daughter Jane gives us the following picture of her father: 'Creative scientific enquiry and rationality were paramount in Jim's approach to life, together with stoicism. He cast a cold eye on anything that seemed motivated by greed, cruelty, self-delusion or sentimentality and had long trained himself to keep emotion under control. As a young man on holiday in pre-war Munich he had seen that city's mayor bundled into a car never to return and had once been surrounded by a huge crowd stirred to frenzy by Adolf Hitler: "Appalling. I could only stand there and wait for them to disperse," he said later. He endured conflicts and tragedies in his personal life by immersing himself even deeper in his work. A natural shyness as well as this guardedness could at times make him seem forbidding but he was deeply attached to his family and friends.' (Jane Robertson, pers. comm. to OM 2004).

As might be expected from this description, when he came in retirement to write a book, it was on common sense in science. In the book, he set out his views on topics from relativity (where he accepted that it was an explanation at the physical level but refuted its biological implications rather as Dr Johnson dealt with Bishop Berkeley; and Rendel always heeded Johnson's sage advice: "Clear your mind of cant") to the future of humanity, from mathematical physics to consciousness. Two of the authors of this memoir read chapters as they were written and had many deeply stimulating discussions over points of disagreement. Characteristically, Rendel would modify what he had written only if he had been convinced that he was wrong; if there was doubt, he left his words to stand. It is a matter of deep regret that he had not found a publisher at the time of his death, for the book embodied profound thought, vast experience and a generous scepticism that are rarely combined in one person. Although not all the references are clear without excessive explication, a letter about the book to one of us gives a little of the flavour of his thinking:

12th July 1996  

75 Falls Road
Wentworth Falls
NSW 2782

Dear Oliver-Bombardment will depend on shape; one does not know how 'pull' will depend on shape. Perhaps the difference has already been fed in to the argument but I can't find an account of the argument.

Metaphysics and semantics get tangled together from time to time. I take Newton to mean that there is some REASON for things coming together, for an arrow leaving a bow etc.: he calls it a force and then relates forces. We can take it for granted that things do come together because they are observed to do so. Why? Not because space is curved.

Particles moving at right angles when struck, yes, thanks, I should have been less precise. With a component at a perpendicular to the path of the photon or some such phrase. Then what happens to the photon.

Common sense. I can see that common sense is not a substitute for the scientific method. The scientific method, call it what you will, is something designed to find out how things fit together; it is not the best way of explaining to the general public what scientists suppose it has discovered-for the time being. Surely as an explanation common sense is OK?

I think Heidegger has got his history wrong. Sometimes he is right but not always. There is a distinction between finding out how to do something by trial & error and then asking why does it go like that and being led by an experiment to see if a technique will work. I can't draw a line between science & technology vis a vis cause and effect.

Many thanks for your comments. I don't follow the one about plants & turbines obeying the same rules according to Newton but not according to me. If true then g is not a push.

Yrs sincerely,
Jim.'

He also produced a short proof, almost short enough to fit in a margin, of Fermat's Last Theorem. This was rather an embarrassment to him because he was well aware that many cranks had produced short proofs containing well- or ill-concealed fallacious reasoning. Two of us could not find defects, but when we showed it to a professional pure mathematician, he identified a line that he considered assumed what had to be proved. Rendel was not happy with this statement, but had still not sent the paper to a journal at the time of his death. There the matter rests, yet one feels that, since the Theorem has been proven correct, a short proof may be waiting to be discovered.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.15, no.2, 2004. It was written by:

• Ian Franklin, Belair, SA.
• Geoff Grigg, Lane Cove, NSW.
• Oliver Mayo, CSIRO Livestock Industries, SA.

Acknowledgments

We thank Sandy Rendel, Tresham Rendel, Jane Robertson, Sandy Robertson, Julia Szulerowski and Polly Woods for assistance with information on JMR's family background, life with JMR and his move to Australia. We thank George Fraser, Bill Sobey, Peter Steele and other colleagues for other assistance in the preparation of this paper.

References

• Australian Academy of Technological Sciences and Engineering 1988 Technology in Australia 1788-1988. Melbourne, Australian Academy of Technological Sciences and Engineering.
• Bagheri-Chaichian, H. and Wagner, G. P. 2002 Evolution of dominance through incidental selection. SFI working papers. 2002: #02-11-064
• Bagheri-Chaichian, H., Hermisson, V., Vaisnys, J. R. and Wagner, G. P. 2002 Epistasis and the kinetics of phenotypic robustness in metabolic pathways SFI working papers. 2002: #02-09-050
• Bennett, J. H. and Binet, F. E. 1956 Association between Mendelian factors with mixed selfing and random mating. Heredity 10 51-55.
• Chalmers, D. J. 1996 The Conscious Mind: In Search of a Fundamental Theory. Oxford University Press.
• Clark, R. W. 1968 J.B.S: The Life and Work of J. B. S. Haldane. London, Hodder & Stoughton.
• Fisher, R. A., and Yates, F. 1963 Statistical Tables for Biological, Agricultural and Medical Research. 6th edn. Edinburgh, Oliver and Boyd.
• Fisher, R. A. 1999 The Genetical Theory of Natural Selection: A Complete Variorum Edition. Edited with an introduction and notes by Henry Bennett. Oxford University Press.
• Gibson, G. and Wagner, G. P. 2000 Canalization in evolutionary genetics: a stabilizing theory? BioEssays 22 372-80.
• Harvey, W. 1657 letter, written six weeks before his death, to Jan Vlakfeld of Haarlem, published in The Works of William Harvey: Translated from the Latin with a Life of the Author by R. Willis. London, Sydenham Society, 1847.
• Kitami, T. and Nadeau, J. H. 2002 Biochemical networking contributes more to genetic buffering in human and mouse metabolic pathways than does gene duplication. Nature Genetics 32 191-4.
• Martin, B. 1997 Suppression Stories. Wollongong, Fund for Intellectual Dissent.
• Mayo, O. 1983 Natural Selection and Its Constraints. London, Academic Press.
• Mayo, O. 2002 Arguments concerning government's investment in research. In Liberty and Research and Development: Science Funding in a Free Society. Edited by Tibor R. Machan. pp. 1-25. Stanford, California, Hoover Institution Press.
• McCann, D. A. and Batterham, P. 1993 Australian genetics: a brief history Genetica 90 81-114.
• Omholt, S. W., Plahte, E., Øyehaug, L. and Xiang, K. 2000 Gene regulatory networks generating the phenomena of additivity, dominance and epistasis. Genetics 156 969-980.
• Rendel, J. M. 1967 Canalisation and Gene Control. London, Logos Press.
• Robertson, A. 1977 Conrad Hal Waddington. Biographical Memoirs of Fellows of the Royal Society. 23 575-622.
• Waddington, C. H. 1957 The strategy of the Genes. London, George Allen & Unwin Ltd.
• Wagner, G.P., Booth, G. and Bagheri-Chaichian, H. 1997 A population genetic theory of canalisation. Evolution 51 329-347.

Publications of J. M. Rendel

1. Rendel, J. M. 1940 Note on the inheritance of yellow bill colour in ducks. Journal of Genetics 40 439-440.
2. Rendel, J. M. 1941 Some factors influencing the weight of table ducklings and the hatchability of ducks' eggs. Empire Journal of Experimental Agriculture 9 50-56.
3. Rendel, J. M. 1944 Symposium 'Application of genetics to plant and animal breeding'. Genetical Society Nature 153 780.
4. Philip, U., Rendel, J. M., Spurway, H. and Haldane, J. B. S. 1944 Genetics and karyology of Drosophila subobscura. Nature 154 260-2.
5. Rendel, J. M. 1944 Genetics and cytology of Drosophila subobscura. II. Normal and selective matings in Drosophila subobscura. Journal of Genetics 46 287-95.
6. Rendel, J. M. 1945 Variations in weights of hatched and unhatched duck's eggs. Biometrika 33 48-58. With appendices by J. B. S. Haldane.
7. Rendel, J. M. and Suley, A. C. E. 1948 Genetics and cytology of Drosophila subobscura. III. Transplantation of eye-buds between Drosophila subobscura and Drosophila melanogaster. Journal of Genetics 49 38-41.
8. Rendel, J. M. 1950 Experimental analysis of the inheritance of productivity and growth in pigs. Animal Breeding Abstracts 18 235-240.
9. Rendel, J. M. and Robertson, A. 1950 Estimation of genetic gain in milk yield by selection in a closed herd of dairy cattle. Journal of Genetics 50 1-8.
10. Robertson, A. and Rendel, J. M. 1950 The use of progeny testing with artificial insemination with dairy cattle. Journal of Genetics 50 21-31.
11. Rendel, J. M. and Robertson, A. 1950 Some aspects of longevity in dairy cows. Empire Journal of Experimental Agriculture 18 69.
12. Rendel, J. M., Robertson, A. and Alim, K. 1951 The extent of selection for milk yield in dairy cattle. Empire Journal of Experimental Agriculture 19 295-301.
13. Rendel, J. M. 1951 Mating of Ebony vestigial and wild type Drosophila melanogaster in light and dark. Evolution 5 226-230.
14. Rendel, J. M. 1952 White Heifer Disease in a herd of dairy Shorthorns. Journal of Genetics 51 89-94.
15. Rendel, J. M. 1953 Heterosis. American Naturalist 87 129-138.
16. Rendel, J. M. 1953 Conclusions from some recent research into animal breeding. Journal of the Australian Institute of Agricultural Science March, pp. 2-7.
17. Rendel, J. M. 1954 Inheritance of birthcoat in a flock of improved Welsh mountain sheep. Australian Journal of Agricultural Research 5 297-304.
18. Rendel, J. M. 1954 The use of regressions to improve heritability. Australian Journal of Biological Sciences 7 368-378.
19. Robertson, A. and Rendel, J. M. 1954 The performance of heifers got by artificial insemination. Journal of Agricultural Science 44 184-207.
20. Rendel, J. M. and Kellerman, G. M. 1955 Deoxyribonucleic acid content of marsupial nuclei. Nature 176 829.
21. Rendel, J. M. 1955 Dwarfism in cattle. The Australian Shorthorn August.
22. Rendel, J. M. and Sheldon, B. L. 1956 Effect of cold treatment on mutation in Drosophila melanogaster. Australian Journal of Agricultural Research 7 566-73.
23. Rendel, J. M. 1956 Cattle breeding for the tropical North. In Beef Cattle in Australia (ed. F. O'Loghlen), pp. 77-83. Sydney, F. H. Johnston Pub. Coy.
24. Rendel, J. M. 1957 Relationship between coincidence and crossing-over in Drosophila. Journal of Genetics 55 95-99.
25. Rendel, J. M., Robertson, A., Asker, A. A., Khishin, S. S. and Ragab, M. T. 1957 The inheritance of milk production characteristics. Journal of Agricultural Science 48 427-432.
26. Rendel, J. M. 1958 The effect of age on the relationship between coincidence and crossing-over in Drosophila melanogaster. Genetics 43 208-14.
27. Rendel, J. M. 1958 Natural and artificial selection. Australian Journal of Science 22 22-27.
28. Rendel, J. M. 1959 Optimum group size in half-sib family selection. Biometrics 15 376-81.
29. Rendel, J. M. 1959 Canalisation of the scute phenotype of Drosophila. Evolution 13 425-39.
30. Rendel, J. M. 1959 Variation and dominance at the scute locus in Drosophila melanogaster. Australian Journal of Biological Sciences 12 524-33.
31. Rendel, J. M. 1960 Animal improvement. Journal of the Australian Institute of Agricultural Science 26 183.
32. Rendel, J. M. and Sheldon, B. L. 1960 Selection for canalization of the scute phenotype in D. melanogaster. Australian Journal of Biological Sciences 13 36-47.
33. Rendel, J. M. 1961 Evolution of dominance. In The Evolution of Living Organisms: A symposium of the Royal Society of Victoria held in Melbourne, December 1959. pp. 102-110.
34. Rendel, J. M. 1961 Consciousness: can it be explained in terms of physics? Australian Scientist 1 149-153.
35. Rendel, J. M. 1962 The relation between gene and phenotype. Journal of Theoretical Biology 2 296-308.
36. Rendel, J. M. 1963 Correlation between the number of scutellar and abdominal bristles in Drosophila melanogaster. Genetics 48 391-408.
37. Moule G. R., Norman M. J. T., Jones R. J. and Rendel J. M. 1963 Development of pastures and beef cattle for northern Australia. UNCSAT, 1963. Pap. no. E/CONF.39/C/396.
38. Rendel, J. M., Sheldon, B. L. and Finlay, D. E. 1964 Effect of homozygosity on developmental stability. Genetics 49 471-84.
39. Rendel, J. M. 1965 Effects of genetic change at different levels. Proc. 16 Int. Congr. Zool. vol. 6. Ideas in Modern Biology. pp. 285-95. New York, Natural History Press.
40. Rendel, J. M. 1965 Scutellar bristles in Drosophila: a comment. Heredity 20 137-8.
41. Rendel, J. M. 1965 Bristle patterns in scute stocks of Drosophila melanogaster. American Naturalist 99 25-32.
42. Rendel, J. M., Sheldon, B. L. and Finlay, D. E. 1965 Canalisation of development of scutellar bristles in Drosophila by control of the scute locus. Genetics 52 1137-51.
43. Rendel, J. M., Sheldon, B. L. and Finlay, D. E. 1966 Selection for canalisation of the scute phenotype. 2. American Naturalist 100 13-31.
44. Rendel, J. M. 1968 Genetic control of a developmental process. In Population Biology and Evolution. (ed. R. C. Lewontin) pp. 47-66. Syracuse, NY, Syracuse University Press.
45. Rendel, J. M. 1968 Control of developmental processes. In Evolution and Environment (ed. E. T. Drake) pp. 341-9. New Haven, Yale University Press.
46. Rendel, J. M. 1969 Model relating gene replicas and gene repression to phenotypic expression and variability. Proceedings of the National Academy of Sciences 64 578-83.
47. Sheldon, B. L., Rendel, J. M. and Finlay, D. E 1969 Possible example of a gene affecting allelic recombination in Drosophila melanogaster. Genetics 63 155-65.
48. Johnston, P. G., Pennycuik, P. R. and Rendel, J. M. 1970 Selection for constancy of expression of the Tabby gene in the mouse. Australian Journal of Biological Sciences 23 1061-6.
49. Rendel, J. M. 1971 Myxomatosis in the Australian rabbit population. Search 2 89-94.
50. Rendel, J. M. 1972. Dairy cattle in hot climates. World Review of Animal Production 8 16-24.
51. Rendel, J. M. 1972 Breeding cattle for the Australian North. World Review of Animal Production 8 48-56.
52. Rendel, J. M. and Binet F. E. 1974 The effect of environment on heritability and predicted selection response: a reply. Heredity 33 106-108.
53. Rendel J. 1974 (Moderator). Round table: adaptability of farm animals to tropical conditions. 1st world congress on genetics applied to livestock production, Madrid, Spain. Vol. 2. Madrid: Editorial Garsi, pp. 211-279.
54. Rendel, J. M. 1975 The utilization and conservation of the world's animal genetic resources. Agriculture and Environment 2 101-19.
55. Rendel, J. M. 1976 Is there a gene regulating the scute locus on the third chromosome of Drosophila melanogaster? Genetics 83 573.
56. Rendel, J. M. 1977 Genetic variance and selection. Proceedings of the 3rd International Congress of the Society for Advanced Breeding Research in Asia and Oceania. Animal breeding papers pp. 20 (ii) 12.
57. Rendel, J. M. 1977 Canalisation in quantitative genetics. Proceedings of the International Conference on Quantitative Genetics, August 16-21, 1976. Ames, Iowa State University Press, pp. 23-28.
58. Pennycuik P. R. and Rendel J. M. 1977 Selection for constancy of score and pattern of secondary vibrissae in Ta/Ta-Ta/Y and Ta/+ mice. Australian Journal of Biological Sciences 30 303-17.
59. Rendel, J. M. and Evans, M. K. 1978 Canalisation of the action of sc' in Drosophila melanogaster. Heredity 41 105-7.
60. Rendel J. M. and Nay T. 1978 Selection for high and low ratio and high and low primary density in Merino sheep. Australian Journal of Agricultural Research 29 1077-86.
61. Rendel, J. M. 1979 Canalisation and selection. In Quantitative Genetic Variation. (eds J. M. Thompson and J. M. Thoday) pp. 139-56. New York, Academic Press.
62. Rendel, J. M. 1980 Low calving rates in Brahman cross cattle. Theoretical and Applied Genetics 58 207-210.
63. Rendel, J. M. 1981 Cattle production in the tropics and improvement through breeding. 32nd Annual Meeting of the European Association for Animal Production, 1981; No. G2.1. 11 pp.
64. Rendel JM 1981 Adaptation of livestock to their environment. Animal genetic resources conservation and management: Proceedings of the FAO/UNEP Technical Consultation, Rome. Food and Agriculture Organization of the United Nations. Pp. 190-200.
65. Rendel, J. M. 1983 Creating new breeds in the wet tropics. Dairy cattle breeding in the humid tropics: Working papers presented at the F.A.O./G.A.O. Expert Consultation held in Hissar, India, February 12-17, 1979. Hissar: Haryana Agricultural University, 1983. pp. 188-198.
66. Rendel, J. M. 1984 Decline in the number of breeds, its consequences and remedies. Genetics: new frontiers. Proceedings of the XV International Congress of Genetics. Volume IV. Applied genetics. New Delhi, Oxford IBH Publishing Co. pp. 23-33.

Notes

1. Myxomatosis, a virus specific to rabbits, the artificial release of which for a time reduced the immense damage done by these animals to the Australian envronment and rural industries.
2. The name Eggatron was a mild play on words, mocking the names of 'big science' equipment such as cyclotron, synchrotron and phytotron.

Professor James Henry ('Jim') Michael 1920–2001 was elected to the Australian Academy of Science in 1973. 

Born in Port Augusta, Jim saw active service during the Second World War. Returning to Adelaide, he completed a PhD in pure mathematics and began a distinguished career as an international expert in mathematical analysis. 

As well as being a mathematician, Jim was a keen golfer and shooter. Jim is remembered as a quiet, gentle man of few words but great integrity. A devoted family man, he is survived by his wife Pat, his daughter Mary Jane, his son Philip and his two grandchildren Ian and Tim Michael.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 24(1), 2013. It was written by M. K. Murray, and D. Parrott.

Jim Morrison was born in Scotland in 1924 and completed his PhD studies in X-ray crystallography at the University of Glasgow before taking up a position with CSIRO in Melbourne in 1949. There he became expert in the construction and operation of mass spectrometers, mainly for the study of ion physics. 

In 1967, he became the Foundation Professor Physical Chemistry at the new La Trobe University in Melbourne where he continued his work in mass spectrometry but was also involved in university leadership that included a period as head of a residential college.

Together with his students, he developed the use of compact, rapid-scanning quadrupole mass spectrometers, linking them in series to allow secondary studies (including photochemistry) of particular ions, but also taking advantage of the speed of the quadrupoles to link them to gas chromatographs for the study of mixtures of organic compounds. In all of this he was a pioneer in the use of computers with spectroscopic instruments. 

Internationally he was a recognised expert, speaking at conferences, establishing collaborations, and spending periods of leave at the University of Utah. After his retirement in 1990 he spent a long period as Emeritus Professor before his death in 2013.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 29(1), 2018. It was written by Ian D. Rae.

Written by T. R. New, C. N. Smithers and A. T. Marshall.

• Introduction
• Early life
• Sudan
• Hong Kong
• Australia
• About this memoir
  • Acknowledgements
  • References
  • Publications

Introduction

Ian Thornton was a fine zoologist, an accomplished academic acknowledged internationally as an authority in his field, and an admired leader and mentor to his colleagues and to generations of students. He came to Australia in early 1968 as Foundation Professor of Zoology at La Trobe University, Melbourne, and remained associated with that department, latterly as Emeritus Professor, for the rest of his life. He guided his department effec­tively until his retirement in 1991 and nurtured standards of excellence in research and teaching, whilst continuing to develop his research interests along two major lines: systematics and biogeography of Psocoptera (an insect order on which he was a recognized world authority) and Pacific-region island biogeography (becoming one of the leading regional biogeographers of his generation). Follow­ing his retirement, Thornton continued and diversified his academic activity, leading further physically strenuous expeditions to Indonesia and Papua New Guinea, publish­ing significant papers and a major book, and fostering international liaisons with universities in Indonesia and Laos.

Early life

Ian Walter Boothroyd Thornton was a Yorkshireman through-and-through, born in Halifax and proud of his heritage and also of the fact that he was born (14 July 1926) on the anniversary of Bastille Day — a fact that somehow infused him in later life with a steadfast, forthright and in part revolutionary outlook, by which he always stood up strongly for what he believed to be just, was a staunch defender of his principles (and of his colleagues and staff), and imbued his dealings at times with an element of fun, and occasional risk. His father John, a Yorkshire dyer who had served in the First World War, died from peritonitis at the early age of 41, when Ian was only 10, but he remained close to his mother Alice Mary, née Crabtree, a Lanca­shire schoolteacher, until she died at the advanced age of 96. He had one sibling, his younger sister Mary Charlotte (later Mary Kitchen). The young Ian was an independ­ent soul. He recollected his early school­days as ‘rather tough’ and that for one period he was caned every morning for transgressions he was going to make that day! Although he was awarded a County Scholarship to Hipperholme Grammar School, he was later sent by his mother after his father’s death to a boarding estab­lishment (Crossley and Porter’s Orphan Home and School), later returning to Hipperholme Grammar when war broke out in 1939. Ian excelled at school, both scholastically, with a number of school prizes to his name, and in sports, being Victor Ludorum in each of his last two years. His competitive nature, so well entrenched during his school years, per­sisted throughout his life. He also always remembered the full details of his paper round, and the hustling skills he acquired on the local church’s snooker table.

Around the end of the Second World War, Ian Thornton served (1944–1948) in the British army. After a short course in military engineering at Birmingham Uni­versity he became an Officer Cadet Sapper in the Royal Engineers. He was later a commissioned officer in the King’s Own Yorkshire Light Infantry and parachutist in the 716 Parachute Brigade Company (6th Airborne Division). He served in the Middle East (Egypt, Palestine and Cyprus, with a trip to India to undertake a course in malaria biology and control), and was demobilized with the effective rank of Lieutenant. In 1948, Ian commenced his studies in Zoology at Leeds University, after marrying Jean (née Jean Frances Brown) at Hipperholme Methodist Church in August of that year. He remembered his undergraduate life fondly, and retained his meticulously transcribed undergraduate course notes throughout his life. Despite never having studied biology at school, Ian had decided in Palestine that he wished to become a zoologist, and the Professor of Zoology at Leeds, Eric Spaul, gave him a chance through a ‘trial year’ (supported also by the Professor of Botany, Irene Manton, whom Ian on their first encounter mistook for the cleaning lady!). He gradu­ated with first class honours in Zoology and Botany, achieved the distinction of Univer­sity Research Scholar (1951) and pro­ceeded to a PhD, supported by a Nature Conservancy Research Studentship and supervised by Edward Broadhead, the authority on British Psocoptera and an ecol­ogist of renown. Psocoptera, small insects that graze on algae and other microflora on the surfaces of bark and foliage, were little known at that time, and Ian was one of the first people to study their ecology in detail. The thesis involved a comparative study of the biology of three coexisting species of Elipsocus on a variety of tree species at Malham Tarn. Field surveys and laboratory experiments were combined elegantly in one of the pioneering studies on psocid ecology that was of much wider relevance in exploring how closely related species could coexist through niche differentiation. This work, completed successfully in 1953, resulted in his first two major papers on psocids (1, 2), and initiated his life-long interest in the twin strands of psocopteran systematics and ecology that would lead eventually to much wider considerations of evolution and biogeography.

Ian then moved from Yorkshire as a fledgling academic to work successively in three very different university environ­ments, on different continents, in each of which his horizons and influence contin­ued to expand. His long-time friend and colleague and fellow Leeds graduate, Alan Marshall, speculated in his eulogy at Ian’s funeral in Melbourne that throughout his life Ian vigorously pursued Charles Darwin’s advice (which Ian had himself quoted in his preface to Darwin’s Islands) that ‘nothing can be more improving to a young naturalist than a journey to distant countries’. Notwithstanding this, the standards and attitudes of a Yorkshire culture were to remain.

Sudan

Ian’s first appointment on leaving Leeds was as Lecturer in Zoology at the then Gordon Memorial College of Khartoum (later to become the University of Sudan [1956] but then affiliated with the Univer­sity of London), for a three-year period, 1953–1956. This resulted in short papers on a variety of taxa and topics (scorpions, sea urchins, moths, succession in papyrus communities [3–6]), so diversifying his broad zoological interests and expertise. His major inspiration seems to have been the hydrologist Julian Rzoska, who was working on the Nile as a biological system and was perhaps instrumental in introduc­ing Ian to ‘big picture ecology’, founded in the study of detail. Psocoptera took a tem­porary ‘back seat’.

Hong Kong

Thornton’s interests in Psocoptera re- established firmly when he moved to the University of Hong Kong as Senior Lec­turer in Zoology (in a department then led by David Barker and later by John Phillips) in 1956. Here he was to remain for the next eleven years. His major initial research thrust was to collect and describe the local psocid fauna (7–11, 14), and to attempt to place them properly in the wider perspec­tive of the fauna of south-east Asia and the western Pacific. He had met Lin Gressitt and other Pacific-region entomologists who were to become long-term friends at the Pacific Science Congress in Bangkok, soon after arriving in Hong Kong. Thus, in addition to descriptions of a substantial number of new taxa, this period also saw development of Ian’s interests in psocid dispersal and distribution, with studies on the wider fauna of the western Pacific, and dispersal mirrored by captures on ships and aircraft in the region (17, 18). Major studies, in part based on examination of the major regional accumulations of speci­mens at the Bishop Museum, Honolulu, resulted in conjunction with his post­graduate students, Wong Siu Kai (Peri­psocidae and Ectopsocidae [26]), Lee Soo-Seong (Pseudocaeciliidae [23]) and Chui Wun Duen (Violet) (Hawaiian and Micronesian taxa); a fourth psocidological student of that era, Woo Kam Tien (Anita) moved with Ian to La Trobe, where she completed her analysis of the Galapagos psocid fauna (33). The results were a much fuller picture of the regional Psocoptera, with strong evolutionary and distributional underpinning to help explain the character­istics of the fauna. As later at La Trobe, Ian also supervised students working on a variety of non-psocid projects in Hong Kong; several of these were focused on the biology of rice pests, in relation to a col­laborative project with Alan Marshall and Cliff Lewis of Imperial College.

These studies on psocids thus laid a solid grounding as stimuli for development of later, more wide-ranging studies. Ian’s interests, eventually to become predomi­nant, in island ecosystems and the pro­cesses of island biogeography were founded in psocids and during the Hong Kong phase of his career. A brief visit to Hawaii in 1961 indicated the explosive speciation of several psocid genera there — rivaling and to some extent paralleling the better documented case of Drosophila on the archipelago. He spent a year at the University of Hawaii’s Institute for Advanced Studies (1963) as a Visiting Senior Scholar, using the time to collect intensively in as many parts of the archipelago as he could reach. The resulting taxonomic monographs (57, 63, 93), mostly not published until some years afterward, are of lasting importance. They changed dramatically the earlier perspective of Hawaiian psocids given in the Insects of Hawaii monograph (Zimmerman 1948).

That year led to seminal changes in Ian Thornton’s thinking and his approach to research. Psocoptera became more firmly tools for exploration of wider evolutionary processes, rather than simply things to be described and enumerated in their own right, although the importance of doing this remained to ensure the reliability of the data he used. His maturing focus on processes of speciation on oceanic archi­pelagos was treated to what was effectively an independent replicate study in 1967, with a three-month stay on the Galapagos Archipelago, the biota of which had so inspired Charles Darwin more than a century before. This study revealed intriguing parallels with Hawaii and also some significant differences. The back­ground information available was far less — simply, no psocids had been recorded previously from any of the Galapagos Islands. The major taxonomic outcome (published jointly with Anita Woo [33]) recorded 39 species, of which 18 were described as new. The similarities and dif­ferences between the faunas of these two archipelagos were important in the evolu­tion of Thornton’s thinking. He regarded the Galapagos fauna as at a much earlier stage of evolution than that of Hawaii.

Hong Kong was not wholly about pso­cids! Ian’s interests encompassed other topics, such as the genetics of the white tigers of Rewa ([27] this resulting from a time he was marooned in Calcutta, thwarted from a planned visit to the Andaman Islands). He also collaborated in the classic studies of the genetics of the mimetic swallowtail butterfly Papilio memnon led by Cyril Clarke and Philip Sheppard, by collecting for them in Pala­wan, Hong Kong and parts of Indonesia (28). A book on Insects of Hong Kong he initiated with Phyllis Hore was later com­pleted by Dennis Hill (book 2). The main impetus for this was Ian’s realization that there was considerable need for a locally focused entomology text, so that his stu­dents did not have to rely on those written more centrally for students in the northern hemisphere. Perhaps the greatest intellec­tual outcome from the Hong Kong years, though, was his widely read book Darwin’s Islands (book 1), mostly written in the year after he left the university and for a decade or more the standard account of the natural history of the Galapagos. The book was translated into Japanese following its initial publication in New York. The wide survey of the islands’ biota is interspersed with numerous personal observations interwoven with the established literature, as well as with ideas on evolution and conservation. As importantly, though, it is immensely readable, and the lucidity and insights that came to characterise Ian’s teaching and writing are already evident.

His academic progress was marked by promotion to Reader in Zoology (October 1966), but Ian had long also played a full part in the corporate life of the university. He was Acting Head or Head of the Depart­ment of Zoology on several occasions and Dean of the Faculty of Science, 1960–1963. He served on many boards and committees (including the University Senate, 1961–1965), and ‘Y. C. W.’ wrote in the University of Hong Kong Gazette (1967) on Ian’s departure, ‘Those of us who have at one time or another sat at the same conference table with him will remember his frankness, his keen observation, and his commonsense approach to problems’. These attitudes persisted, as did his con­cerns for students and colleagues. Again from Y. C. W.: ‘Dr Thornton enjoys a high reputation as a teacher and has a genuine concern for all his students…. Many of his students and junior colleagues will not forget the help and guidance they received from him on academic and other matters.’ Indeed, throughout his career, Ian was an inspirational teacher. In Hong Kong he was instrumental in introducing highly ‘urban­ized’ Chinese students to rigorous field work on the then remote Lantoa Island. He consistently trusted his judgement of students, even to the extent of confronting eminent external examiners when he considered their opinions deficient. His tabletop duel (using toy swords, and ending with both protagonists falling off) with J. Z. Young (University College, London) resulting from one such defence of his students’ marks has passed into folklore.

Australia

Ian and Jean, with their children Angus and Jane, arrived in Australia on the Royal Interocean Lines ship ‘Tjiluwah’ on 6 January 1968, which he recalled as a ‘100 degree day’, to take up the Founda­tion Chair of Zoology at La Trobe Univer­sity, Melbourne, Ian having selected this from amongst the three chairs he was offered around that time. The late 1960s was an exciting time in Australian univer­sities, with an air of optimism brought about by the establishment of several new institutions, amongst which the promise of La Trobe was influential in Ian’s decision. Although interview (by a committee including Macfarlane Burnet) and selec­tion were rigorous, Ian recorded that he was subsequently first offered the job by the then Vice-Chancellor, David Myers, whilst they occupied adjacent urinal stalls in the gentleman’s toilet! He formally accepted a few days later. Ian set about establishing a Zoology Department (ini­tially as a non-departmentalized part of a wider School of Biological Sciences, fol­lowing the educational philosophy of the university’s founders), based on his belief, from which he never deviated, that ‘Zool­ogy is the study of animals, not just of books about animals’. He recognized the need to recruit colleagues, predominantly focused on ‘whole animal biology’, who were capable of communicating both knowledge and enthusiasm to their stu­dents. The major thrust of the department was to be ‘terrestrial zoology’; at that time Monash University (although also strong in terrestrial zoology) had firmly estab­lished regional leadership in freshwater biology (through Bill Williams and his colleagues), and the logistic difficulties of developing a strong marine programme were formidable. Ian also believed that the Professor should be the Head of Depart­ment, as both academic leader and mentor, and he fulfilled both roles for as long as he was allowed to do so (that is, until his retirement in 1991).

The scope of psocid studies initiated whilst in Hong Kong continued as the major research focus of Thornton’s first decade in Australia, but became conceptu­ally expanded and geographically concen­trated on the biogeography of the Pacific region, particularly the western side, and in particular on the psocid faunas of the Melanesian arcs of islands. Much of this work was based on Ian’s own field expedi­tions in a long-term ARGC-funded project (1971–1982), with Courtenay Smithers and (to a far lesser extent) Tim New as collaborators. Thus, over a period of some twenty years from the mid-1960s, Ian visited and collected psocids in Sri Lanka, the Himalayan foothills, Malaysia, Japan, many parts of Indonesia, Papua New Guinea, the Solomon islands, Palawan, many parts of the Melanesian arcs includ­ing Norfolk Island, New Zealand, the New Hebrides, Fiji, Tonga, the Society Islands, the Galapagos, Mexico, Colombia, Ecuador, Peru, Chile (including the Juan Fernandez archipelago), Argentina and Hawaii. He maintained detailed field jour­nals for most of his field work, and the progressive list of places visited reads like a major gazeteer for this vast region. These studies gave him a unique personal per­spective on an insect order and its evolu­tion over a substantial part of the world.

Most of the field work was undertaken on shoestring budgets, and Ian’s Yorkshire upbringing and philosophy rendered him reluctant to operate on anything more than restricted personal financial input. Wher­ever possible, he would bargain hard to reduce costs of accommodation, car hire and so on, and he cared little what he ate — a packet of cornflakes was just as satisfying as a three-course spread. However, despite strenuous physical activity, he rarely suc­cumbed to gastric or other upsets in the field. Much of the collecting was in remote areas, with a general tendency to move upward from lowlands to mountains on the basis that these would yield more ‘typical’ or endemic psocids than the more disturbed lower regions. In part, this was common sense in facilitating access to less-disturbed habitats. Ian frequently turned his eyes, and his body, toward the hills. Distributions along altitudinal gradients intrigued him, and he sometimes claimed that simply climbing up Mt Rinjani (Lombok) and seeing the changes along the way was a very fine lesson on tropical biology for any student to undertake.

Collecting trips with Ian tended not to be luxurious and relaxing, despite the envious comments made by colleagues who did not participate and thought of New Guinea and like places as ‘romantic’. He worked hard, remained focused on his objectives, and maintained a positive atti­tude under sometimes appalling and dan­gerous conditions. Ian’s competitive nature persisted on field trips so that visits to the local ‘expatriate club’ in (for example) remote parts of New Ireland or New Britain could become ‘interesting’. He prided himself on the skills at snooker obtained in his youth and commonly challenged the local champion to a game, which he resolved to win. If things were not going his way (a common occurrence, simply because most such local players sometimes seemed never to move away from the table!), a frequent gambit was to pause and casually ask his opponent whether the ball he was about to address was of a particular colour. After the usual surprised/annoyed retort, Ian would point out (correctly) that he was colour-blind and, having so discon­certed his opponent, commonly went on to win the game. Ian’s persistence, neverthe­less, took him and his collecting compan­ions to many remote areas that had never been explored before, some of which have now been changed dramatically by human pressures. The taxonomic treatment and faunal analysis of the accumulated Psocop­tera added massively to knowledge of this complex and rapidly changing region. The succession of descriptive papers, many of them co-authored, and illustrated by Justine O’Regan, Jodie Kernutt, John Greer, Jenny Browning or Tracey Carpenter, are signifi­cant additions to the psocid literature. Alto­gether, Ian (alone or with his co-authors) described almost 750 new species of Pso­coptera, a significant proportion of the doc­umented world fauna. His work on psocids was recognized by the small global frater­nity of psocidologists in dedicating eight species to him (as named ‘thorntoni’) and in the genus Thorntoniella whilst he was alive; a commemorative volume of papers on Psocoptera (Garcia Aldrete et al. 2005) augments these by a further two genera (Ianthorntonia, Thorntonodes) and four species.

However, this basic taxonomic work was simply a template for Ian’s increasing interests in island biogeography and pat­terns of distribution and speciation. Sub­stantial papers on the distribution and origins both of taxa (e.g. Philotarsidae) and faunas (Hawai’i) are classics of much wider interest than to psocidologists alone. Ian’s DSc degree (Leeds, 1984) recognized the importance of this documentation and his developing syntheses, which later came to constitute some of his most significant work and to establish him among the forefront of modern Pacific-region biogeographers.

The second major theme, developed from the early 1980s on, was to lead to what many peers regard as Ian’s finest academic achievements. In 1982, with Ann (he had married Ann Juliana Patterson in 1980, following the dissolution of his first marriage in the mid-1970s) Ian had his first sight of the area that was to become his major scientific passion for the next decade and more — the Krakatau islands, nestled in the Sunda Strait between Sumatra and Java. Ian later noted his recur­ring feelings of excitement and awe each time the small fishing boats used for travel to the islands entered the caldera and moved along the base of the imposing towering sheer cliff face (800 m high) of Rakata. On that initial visit, Ian recognized the unique opportunity Krakatau provided for studying the colonization processes and development of tropical communities and ecosystems from a tabula rasa begin­ning. In contrast to Hawaii, where the emphasis of his studies had been on post- colonization radiations related to isolation, his perspective now broadened further to consider and study the initiation and devel­opment of tropical communities. The unique natural laboratory of the Krakataus comprised two distinct temporal sequences for studying the development of tropical systems. First, the cataclysmic 1883 eruption is widely believed to have obliter­ated all life from the islands, so that the condition of vegetation and animal assem­blages on the three older islands (Rakata, Sertung, Panjang) in the 1980s represented the outcomes of a century of re-establish­ment from source areas of Java and Sumatra, each more than 40 km away. Second, and nested within this, the island of Anak Krakatau (‘Child of Krakatau’) emerged lastingly from the sea in the centre of the caldera in 1930, undoubtedly virgin land and providing a second, much younger sequence for study of colonization from the much closer source areas of the other three islands. Six expeditions to the Krakataus were organized and led by Ian, extending over almost a decade from 1984. They involved many colleagues and col­laborators, and led to a series of papers of lasting interest and relevance in island biogeography. They culminated in Ian’s magnum opus Krakatau: The Destruction and Reassembly of an Island Ecosystem in 1996 (book 3), a book widely regarded as Ian’s finest academic achievement, and recognized by winning the 1996 Profes­sional Scholarly Publications Award of the Association of American Publishers, Bio­logical Sciences Category. These expedi­tions were hard work, at times frustrating, but probably all participants (including several honours and graduate students working in the tropics for the first time) viewed them as highlights in their aca­demic careers. Camping on Anak Krakatau provided a remote but idyllic scenario in that harsh environment, and it came to be a place that Ian (and his companions) loved — despite the ever-present threat of vol­canic activity, the fact that all food and water had to be carried to the island from Java, and the undoubted terrors of unpre­dictable sea crossings in rough weather. Tim New went on four of those expeditions and noted that Ian’s participation was enthusiastic and dynamic. Indonesian counterpart scientists became lasting friends, and sojourns in Bogor or else­where during the lengthy process of obtaining permits and other documentation allowed opportunity to collect in a variety of possible source areas for the Krakatau fauna. As in much of Ian’s earlier work, psocids were a focal group but now only one of numerous biota (even including bacteria [73–77] and soil nematodes) incorporated into the emerging picture. A highlight for Australian expeditioners was the participation of scientists from many other parts of the world. With the closest parallel study to the one on Anak Krakatau being based on the emergence of Surtsey (off Iceland, and which Ian had visited a year or so previously), a brief visit to Anak in 1990 by Sturla Fridriksson helped to foster insights from a broad and authorita­tive base, and provided an opportunity to discuss parallels more closely.

The major importance of the ‘Krakatau study’ was recognized by Ian being awarded the John Lewis Gold Medal by the Royal Geographical Society of Australasia (1992) and his election to the Fellowship of the Australian Academy of Science (1995). Later, Ian turned his attention to even wider aspects of the colonization of volcanic islands, appraising the role of Sebesi in the northern part of the Sunda Strait as a ‘stepping stone’ for colonization of the Krakataus (119), and (then in his 70s) making a further physically strenuous expedition to explore Mot Mot, a rare example of an island in a lake in the closed caldera of a volcanic island (Long Island) in Papua New Guinea. His major collabo­rator on this exploit was John Edwards, who had also visited Krakatau with Ian and who had worked extensively and innovatively on the colonization patterns following the eruption of Mt St Helens, Washington State, USA, in 1980. And, as for the Krakatau studies, Ian edited the series of papers to ensure that they appeared in co-ordinated and accessible form rather than being scattered widely. The Long Island papers constituted a special issue of the Journal of Biogeography, following the earlier Krakatau papers grouped in Philosophical Transactions of the Royal Society of London (1988, 1990) and GeoJournal (1992, this last being the proceedings of a two-day session organized by Ian at a Pacific Science Congress in Honolulu).

Ian’s zest for life and for his science was infectious. He revelled in academic dis­course and persisted with argument until he got responses that (at least for the time being) satisfied his curiosity. He was a perceptive reviewer of grant applications and manuscripts and served on several editorial boards, as well as a term as Vice- President of the Australian Entomological Society. The example of enthusiasm and wonder he set to his students and col­leagues is a lasting one, and generations of undergraduates had their scientific atti­tudes and perceptions honed and focused by his influence. Ian enjoyed teaching, both in the formal lecture-theatre context and on field courses, where his staying power was legendary. New was usually among the earliest risers on such trips, and recalls that it was common to find Ian still ‘instructing’ (not necessarily solely on scientific topics!) at around 5 a.m., with his youthful undergraduate audience ever more aware that they were due to start a strenuous day of field work within a couple of hours….

In his early days at La Trobe, Ian was a key instigator in the formation of the School of Biological Sciences. He was a natural leader who inspired loyalty in his staff. His colleagues agreed with his strong belief that, as Professor, he should lead his department, and at the time when most departments at La Trobe were encouraged to elect their head Ian was endorsed as ‘permanent chairman’. He fought hard to defend the concept that a University should be a community of scholars free to pursue their research interests without interference from government. He strongly resented the rise of cohorts of ‘academis­trators’ (his term, not entirely complimen­tary), and on occasion urged academic disobedience to resist externally imposed changes. He wrote formally to the Univer­sity Council in 1988 under the heading ‘Take up the Mace!’ (a reference to the University’s ceremonial mace carried on formal academic occasions but — as far as we know — never used in anger), ‘asking Council for its support in the defence of my rights and responsibilities as a profes­sor, and in defence of my discipline from outside interference’. Elsewhere, he argued his belief that ‘No-one realizes that universities cannot be run like businesses, because good universities are inherently inefficient operations — decisions are questioned, considered, mulled over, in a collegiate system’. The then recent changes to university priorities in Australia depressed him greatly, not least because he saw the opportunities for young people being eroded as funding and teaching capability declined, to the detriment of Australia’s future. He strongly resented his enforced retirement on grounds of age when he reached 65 and characteristically fought hard against this — even seeking professional advice as to whether he was subsequently able to apply for the job of his replacement, by which time the manda­tory age retirement no longer existed! Ian served three periods as Dean of Biological Sciences (1970–1972, 1979–1981, 1985–1987) and was Acting Vice-Chancel­lor on two occasions. He sat on most of the University’s major boards and committees, where his determination, humour and abil­ities to think rapidly and laterally about many complex issues were useful counters to the tedium that some such bodies can adopt, and gained him the respect of col­leagues throughout the institution. The University recognized his contributions, in conjunction with his scientific stature, by the posthumous award of the DSc degree, honoris causa, coincidentally presented to Ann on the first anniversary of his funeral.

Post-retirement, Ian lectured for many years in Natural Resource Management to Applied Sciences students at the Holmes­glen College of TAFE, inspiring several of them to move on to university studies in related fields. In addition to his scientific achievements, Ian had a strong interest and involvement in fostering Australian/Indo­nesian academic co-operation and educa­tional development. In the years after retirement, he was an academic adviser or guest lecturer at Udayana University (Bali), Mataram University (Lombok) and other Indonesian universities. As an Indo­nesian colleague recently expressed it, ‘Ian Thornton showed how Australian and Indo­nesian colleagues could work together’. His death occurred in Bangkok whilst he was returning from Laos, where he was advising the National University on the implementation of basic science courses. He is survived by Ann and three stepchil­dren, two adopted children from his earlier marriage to Jean, and six grandchildren.

Many zoologists have made notable con­tributions to different fields within their discipline, but Ian Thornton is memorable for the number of very different fields to which he made highly significant contri­butions. This stemmed from his immutable belief that a scholar should be allowed to follow his interests, and to his own acumen when interesting opportunities arose. He never took short cuts —a casual query from a student could engage him for several hours. Once he had decided a particular course, if the nature of the progress demanded laborious enterprise or even dan­gerous fieldwork he would not be deflected. He adhered firmly, and in our opinion cor­rectly, to the ideal that zoogeography, ecology and indeed any aspect of the evolu­tion of a group can be understood fully only after adequate systematic study. The consid­erable sacrifice and efforts needed to pursue fieldwork to collect Psocoptera in remote areas were simply ‘part of the game’ that he played so ably over much of his academic life.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.16, no.1, 2005. It was written by:

• T. R. New, Department of Zoology, La Trobe University, Victoria
• C. N. Smithers, The Australian Museum, Sydney
• A. T. Marshall, Department of Zoology, La Trobe University, Victoria

Acknowledgments

We appreciate greatly the considerable help and advice given by Ann Thornton for this memoir. Longstanding colleagues and friends at La Trobe, particularly Pat Woolley, also generously shared their thoughts and reminiscences with us. The photograph shows Ian Thornton on Anak Krakatau, with the cliff of Rakata in the background.

References

• Garcia Aldrete, A.N., Lienhard, C. and Mockford, E.L. (2005). Thorntonia. A Com­memorative Volume for Ian W. B. Thornton. Publicaciones Especiales 20. Insituto de Bio­logia, Universidad Nacional Autonoma de Mexico. 205 pp.
• Zimmerman, E.C. (1948). Insects of Hawaii. Vol. 2. University of Hawaii Press, Honolulu.

Publications

Books

1. Thornton, Ian. (1971). Darwin’s Islands: a Natural History of the Galápagos. Double­day, Natural History Press, New York. 322 pp.
2. Hill, D.S., Hore, P. and Thornton, I.W.B. (1982). Insects of Hong Kong. Hong Kong University Press, Hong Kong. 503 pp.
3. Thornton, Ian. (1996). Krakatau – The Destruction and Reassembly of an Island Ecosystem. Harvard University Press, Cam­bridge, Mass., USA. 346 pp.

Chapters of books

1. Thornton, I.W.B. (1991). ‘Krakatau – studies on the origin and development of a fauna’, in The Unity of Evolutionary Biology (Proceed­ings of the Fourth International Congress of Systematic and Evolutionary Biology), E.C. Dudley (Ed.), pp. 396–408. (Diosco­rides Press, Portland, USA.)
2. Thornton, I.W.B. (1996). ‘The origins and development of island biotas as illustrated by Krakatau’, in The Origin and Evolution of Island Biotas. New Guinea to Eastern Poly­nesia: Patterns and Processes, A. Keast and S.E. Miller (Eds), pp. 67–90. (SPB Academic Publishing, The Netherlands.)
3. Thornton, I.W.B. (1999). ‘The ecology of volcanoes: recovery and reassembly of living communities’, in Encyclopedia of Volcanoes, H. Sigurdsson (Ed.), 1057–1081. (Academic Press, San Diego, USA.)

Edited volume

1. Thornton, I.W.B. (Scientific Ed.) (1992). Krakatau: a Century of Change. GeoJournal 28(2), 83–304 (editorial comments pp. 84–86, 129, 173–174, 232, 292).

Research papers in scientific journals

1. Thornton, I.W.B. and Broadhead, E. (1954). The British species of Elipsocus Hagen (Cor­rodentia, Mesopsocidae). Journal of the Soci­ety for British Entomology 5(2), 47–64.
2. Broadhead, E. and Thornton, I.W.B. (1955). An ecological study of three closely related psocid species. Oikos 6(1), 1–50.
3. Thornton, I.W.B. (1956). Diurnal migrations of the echinoid Diadema setosum (Leske). British Journal of Animal Behaviour 4(4), 143–146.
4. Thornton, I.W.B. (1956). Notes on the biol­ogy of Leiurus quinquestriatus (H. & E. 1829) (Scorpiones, Buthidae). British Jour­nal of Animal Behaviour 4(3), 92–93.
5. Thornton, I.W.B. (1957). Faunal succession in umbels of Cyperus papyrus L. on the Upper White Nile. Proceedings of the Royal Entomological Society of London A 32, 119–131.
6. Thornton, I.W.B. (1957). Notes on the ecol­ogy of the Acacia bagworm, Auchmophila kordofensis (Lepidoptera, Psychidae), in the environs of Khartoum. Sudan Notes and Records 38, 147–150.
7. Thornton, I.W.B. (1959). A new genus of Philotarsidae (Corrodentia) and new species of this and related families from Hong Kong. Transactions of the Royal Entomological Society of London 111(2), 331–349.
8. Thornton, I.W.B. (1959). New species of Peripsocus Hagen 1866 (Corrodentia, Peri­psocidae) from Hong Kong Island, with fur­ther descriptions of Peripsocus similis Enderlein (1903) and Peripsocus quercicola Enderlein (1906). Proceedings of the Royal Entomological Society of London B 28, 37–48.
9. Thornton, I.W.B. (1960). New Psocidae and an aberrant new myopsocid (Psocoptera) from Hong Kong. Transactions of the Royal Entomological Society of London 112(10), 239–261.
10. Thornton, I.W.B. (1961). Comments on the geographical distribution of Pseudocaecilius elutus Enderlein (Psocoptera), with descrip­tions of related new species from Hong Kong. Proceedings of the Royal Entomological Society of London B 30, 141–152.
11. Thornton, I.W.B. (1961). The Trichade­notecnum group (Psocoptera: Psocidae) in Hong Kong, with descriptions of new species. Transactions of the Royal Entomo­logical Society of London 113(1), 1–24.
12. Thornton, I.W.B. (1962). Note on the geni­talia of two New Zealand philotarsids (Insecta: Psocoptera). Journal of the New Zealand Institute of Science 5(2), 241–245.
13. Thornton, I.W.B. (1962). Psocids (Psoco­ptera) from the Batu Caves, Malaya. Pacific Insects 4(2), 441–455.
14. Thornton, I.W.B. (1962). The Peripsocidae (Psocoptera) of Hong Kong. Transactions of the Royal Entomological Society of London 114(9), 285–315.
15. Marshall, A.T. and Thornton, I.W.B. (1963). Micromalthus (Coleoptera: Micromalthidae) in Hong Kong. Pacific Insects 5(4), 715–720.
16. Thornton, I.W.B. (1963). The ecology of closely related species. Proceedings of the 9th Pacific Science Congress, Bangkok, 19–25. (Delivered in 1957.)
17. Thornton, I.W.B. (1964). Airborne Psoco­ptera trapped on ships and aircraft. Pacific Insects 6(2), 285–291.
18. Thornton, I.W.B. and Harrell, J.C. (1965). Air-borne Psocoptera trapped on ships and aircraft, 2 – Pacific ship trappings 1963–64. Pacific Insects 7(4), 700–702.
19. Thornton, I.W.B. (1965). Distribution pat­terns of endemic psocids (Psocoptera) in the Hawaiian Islands. Proceedings of the XIIth International Congress of Entomology, Lon­don 1964, 442–443.
20. Wong, S.K. and Thornton, I.W.B. (1966). Chromosome numbers of some psocid gen­era. Nature 211(45), 214–215.
21. Thornton, I.W.B. (1966). Isolation within archipelagos. Proceedings of the 11th Pacific Science Congress, Tokyo 5, 12.
22. Thornton, I.W.B. and Wong S.K. (1966). Some Psocoptera from West Bengal, India. Transactions of the Royal Entomological Society of London 118(1), 1–21.
23. Lee, S.S. and Thornton, I.W.B. (1967). The family Pseudocaeciliidae (Psocoptera) – a reappraisal based on the discovery of new Oriental and Pacific species. Pacific Insects Monographs 16, 1–116.
24. Thornton, I.W.B. (1967). The measurement of isolation on archipelagos, and its relation to insular faunal size and endemism. Evolu­tion 21(4), 842–849.
25. Thornton, I.W.B. (1967). Wing reduction in endemic Hawaiian psocids. Journal of Natural History 1, 149–150.
26. Thornton, I.W.B. and Wong, S.K. (1967). A numerical taxonomic analysis of the Perip­socidae of the Oriental Region and the Pacific Basin. Systematic Zoology 16(3), 217–240.
27. Thornton, I.W.B., Yeung, K.K. and Sankhalz, K.S. (1967). The genetics of the white tigers of Rewa. Journal of Zoology 152, 127–135.
28. Clarke, C.A., Sheppard, P.M. and Thornton, I.W.B. (1968). The genetics of the mimetic butterfly Papilio memnon L. Philosophical Transactions of the Royal Society of London B 254, 37–89.
29. Thornton, I.W.B. and Wong, S.K. (1968). The peripsocid fauna (Psocoptera) of the Oriental Region and the Pacific. Pacific Insects Mono­graphs 19, 1–158.
30. Wong, S.K. and Thornton, I.W.B. (1968). The internal morphology of the reproductive sys­tems of some psocid species. Proceedings of the Royal Entomological Society of London A 43(1–3), 1–12.
31. Chui, W.D. and Thornton, I.W.B. (1972). A numerical taxonomic study of the endemic Ptycta species of the Hawaiian Islands (Psocoptera: Psocidae). Systematic Zoology 21(1), 7–22.
32. Thornton, I.W.B., Lee, S.S. and Chui, W.D. (1972). Psocoptera. Insects of Micronesia 8(4), 45–144.
33. Thornton, I.W.B. and Woo, A.K.T. (1973). Psocoptera of the Galapagos Islands. Pacific Insects 15(1), 1–58.
34. Smithers, C.N. and Thornton I.W.B. (1973). The Psilopsocidae (Psocoptera) of New Guinea. Proceedings of the Linnean Society of New South Wales 98(2), 98–103.
35. Smithers, C.N. and Thornton I.W.B. (1974). The Myopsocidae (Psocoptera) of New Guinea and New Caledonia. Transactions of the Royal Entomological Society of London 126(1), 91–127.
36. Smithers, C.N. and Thornton, I.W.B. (1974). The Psocoptera of Norfolk Island. Records of the Australian Museum 29(8), 209–234.
37. Thornton, I.W.B. and Smithers, C.N. (1974). The Philotarsidae (Psocoptera) of New Cale­donia. Pacific Insects 16(2–3), 177–243.
38. New, T.R. and Thornton, I.W.B. (1975). Psocomorpha (Psocoptera) collected on recent expeditions to South America. Journal of Entomology B 44(1), 27–80.
39. Smithers, C.N. and Thornton, I.W.B. (1975). The first record of Stenopsocidae (Psoco­ptera) from New Guinea with descriptions of new species. Proceedings of the Linnean Society of New South Wales 100(2), 156–166.
40. Smithers, C.N. and Thornton, I.W.B. (1975). The Psocoptera of Lord Howe Island. Records of the Australian Museum 29(16), 453–471.
41. Thornton, I.W.B., Marshall, A.T., Kwan, W.H. and MA, Q. (1975). Studies on lepido­pterous pests of rice crops in Hong Kong, with particular reference to the Yellow Stem- borer, Tryporyza incertulas (Walk.). Pest Articles and News Summaries 21, 239–252.
42. New, T.R. and Thornton, I.W.B. (1975). Psocomorpha (Psocoptera) from the Malayan Peninsula, including collections from forest canopy. Oriental Insects 9(4), 375–418.
43. Smithers, C.N. and Thornton, I.W.B. (1977). A new genus and some new species of Epi­psocidae (Psocoptera) from the Melanesian Arc. Proceedings of the Linnean Society of New South Wales 102(2), 60–75.
44. Thornton, I.W.B. and New, T.R. (1977). Philotarsidae (Psocoptera) of the Bismarck Archipelago. Pacific Insects 17(4), 451–457.
45. Thornton, I.W.B. and New, T.R. (1977). The Philotarsidae (Psocoptera) of Australia. Aus­tralian Journal of Zoology, Supplementary Series 54, 1–62.
46. Thornton, I.W.B. and Smithers, C.N. (1977). Philotarsidae (Psocoptera) of New Guinea. Pacific Insects 17(4), 419–450.
47. Thornton, I.W.B., Wong, S.K. and Smithers, C.N. (1977). The Philotarsidae (Psocoptera) of New Zealand and islands of the New Zea­land plateau. Pacific Insects 17(2–3), 197–228.
48. Thornton, I.W.B. (1978). White tiger genetics – further evidence. Journal of Zoology, Lon­don 185, 389–394. (Review in Science Report, The Times, 1.ix.78, p.14.)
49. Thornton, I.W.B. and Lyall, I. (1978). Psoco­ptera from Chilean Nothofagus. Pacific Insects 19(1–2), 1–16.
50. Thornton, I.W.B. and Smithers, C.N. (1978). Philotarsidae (Psocoptera) of the Solomon Archipelago. Pacific Insects 18(3–4), 227–233.
51. Smithers, C.N. and Thornton, I.W.B. (1979). Psilopsocidae and Myopsocidae (Psocoptera) of the Bismarck, Solomon and New Hebrides archipelagos. Records of the Australian Museum 32(16), 513–545.
52. Thornton, I.W.B. (1980). Plate tectonics and the distribution of the insect family Philo­tarsidae (Order Psocoptera) in the Southwest Pacific. Palaeogeography, Palaeoclimat­ology, Palaeoecology 31, 251–266.
53. New, T.R. and Thornton, I.W.B. (1981). Psocoptera from central and southern Chile. Pacific Insects Monographs 37, 136–178.
54. Thornton, I.W.B. (1981). Psocoptera of the Fiji Islands. Pacific Insects Monographs 37, 1–105.
55. Thornton, I.W.B. (1981). Psocoptera of the Tongan Archipelago. Pacific Insects Mono­graphs 37, 106–135.
56. Thornton, I.W.B. and New, T.R. (1981). Psocoptera from Robinson Crusoe Island, Juan Fernandez Archipelago. Pacific Insects Monographs 37, 179–191.
57. Thornton, I.W.B. (1981). Psocoptera of the Hawaiian Islands. Parts I and II. Introduction and the nonendemic fauna. Pacific Insects 23(1–2), 1–49.
58. Thornton, I.W.B. (1981). The systematics, phylogeny and biogeography of the psoco­pteran family Philotarsidae. Systematic Ento­mology 6(4), 413–452.
59. Smithers, C.N. and Thornton, I.W.B. (1981). The Psocidae (Insecta: Psocoptera) of New Guinea, including a new coleopteriform genus from high on Mt Wilhelm. Australian Journal of Zoology 29(6), 921–969.
60. Smithers, C.N. and Thornton, I.W.B. (1981). ‘The role of New Guinea in the evolution and biogeography of some families of psoco­pteran insects’, in J.L. Gressitt (Ed.) Biogeo­graphy and Ecology of New Guinea Vol. 2, 621–638 (W. Junk, Publishers, The Hague).
61. Thornton, I.W.B. (Scientific Ed.). (1983). Proceedings of Symposium on Biogeography and Plate Tectonics in the SW Pacific, 15th Pacific Science Congress, Dunedin, (1983). GeoJournal 7(6), 479–564.
62. Thornton, I.W.B. (1983). ‘Vicariance and dis­persal: confrontation or compatibility?’, in I.W.B. Thornton (Ed.) Symposium on Bio­geography and Plate Tectonics in the Pacific. GeoJournal 7(6), 557–564.
63. Thornton, I.W.B. (1984). Psocoptera of the Hawaiian Islands. Part III. The endemic Ptycta complex of species: systematics, distribution and possible phylogeny. Inter­national Journal of Entomology 26(1–2), 1–128.
64. Thornton, I.W.B. (1984). Review of ‘Taxon­omy, phylogeny and biogeography of the genus Cosmopsaltria, with remarks on the historic biogeography of the subtribe Cosmo­psaltriaria (Homoptera: Cicadidae)’ by J.P. Duffels. International Journal of Entomology 26(1–2), 171–173.
65. Thornton, I.W.B. and Smithers, C.N. (1984). The systematics of the Calopsocidae, an Oriental and Melanesian family of Psoco­ptera. Systematic Entomology 9(2), 183–244.
66. Thornton, I.W.B. (1984). Krakatau – the development and repair of a tropical eco­system. Ambio 13(4), 216–225.
67. Thornton, I.W.B. (1984). Psocoptera and Wallace’s Line: collections from the islands of Bali and Lombok. Treubia 29(2), 83–177.
68. Thornton, I.W.B. (1984). An unusual psoco­pteran from New Guinea, and its relation­ships within the Philotarsidae. International Journal of Entomology 26(4), 378–385.
69. Thornton, I.W.B. (1985). The geographical and ecological distribution of arboreal Psoco­ptera. Annual Review of Entomology 30, 175–196.
70. Thornton, I.W.B. (1985). A preliminary sur­vey of the psocopteran fauna of the Krakatau Islands. Proceedings of the Symposium on 100 years development of Krakatau and sur­roundings. Jakarta, L.I.P.I. 466–470.
71. Thornton, I.W.B., Zann, R.A., Rawlinson, P.A., Tidemann, C.R., Adikerana, A.S. and Widjoya, A.H.T. (1988). Colonization of the Krakatau Islands by vertebrates: equilibrium, succession and possible delayed extinction. Proceedings of the National Academy of Science of the USA (Ecology) 85, 515–518.
72. Thornton, I.W.B. and Rosengren, N.J. (1988). Zoological Expeditions to the Krakatau islands, 1984–1985: General Introduction. Philosophical Transactions of the Royal Society of London B 322, 273–316.
73. Graves, S.R., Plummer, D.C., Hives, N., Harvey, K.J. and Thornton I.W.B. (1988). Antibiotic resistance patterns of soil bacteria (Gram-negative rods) from the Krakatau Is (Rakata) and W. Java. Philosophical Trans­actions of the Royal Society of London B 322, 317–326.
74. Graves, S.R., Rosengren, N.J., Kennelly- Merrit, S.A., Harvey, K.J. and Thornton, I.W.B. (1988). Antibiotic resistance patterns and relative concentrations of bacteria (Gram-negative rods) from ash deposits of various ages on the Krakatau Is. Philo­sophical Transactions of the Royal Society of London B 322, 327–338.
75. Graves, S.R., Kennelly-Merrit, S.A., Tidemann, C.R., Rawlinson, P.A., Harvey, K.J. and Thornton, I.W.B. (1988). Antibiotic- resistance patterns of enteric bacteria of wild mammals on the Krakatau Is and W. Java. Philosophical Transactions of the Royal Society of London B 322, 339–354.
76. Graves, S.R., Rawlinson, P.A., Kennelly- Merrit, S.A., McLaren, D.A., Harvey, K.J. and Thornton, I.W.B. (1988). Enteric bacteria of reptiles on Java and the Krakatau Islands. Philosophical Transactions of the Royal Society of London B 322, 355–362.
77. Thornton, I.W.B. and Graves, S.R. (1988). Colonization of the Krakataus by bacteria and the development of antibiotic resistance. Philosophical Transactions of the Royal Society of London B 322, 363–368.
78. Thornton, I.W.B., New, T.R. and Vaughan, P.J. (1988). Colonization of the Krakatau Islands by Psocoptera. Philosophical Transactions of the Royal Society of London B 322, 427–444.
79. New, T.R., Bush, M., Thornton, I.W.B. and Sudarman, H.K. (1988). The butterfly fauna of the Krakatau Islands after a century of colonization. Philosophical Transactions of the Royal Society of London B 322, 445–458.
80. Compton, S.G., Thornton, I.W.B., New, T.R., and Underhill, L. (1988). The colonization of the Krakatau Islands by fig wasps and other chalcids (Hymenoptera, Chalcidoidea). Philosophical Transactions of the Royal Society of London B 322, 459–470.
81. Thornton, I.W.B., New, T.R., McLaren, D.A., Sudarman, H.K. and Vaughan, P.J. (1988). Airborne arthropod fall-out on Anak Kraka­tau and a possible pre-vegetation pioneer community. Philosophical Transactions of the Royal Society of London B 322, 471–480.
82. New, T.R. and Thornton, I.W.B. (1988). A pre-vegetation population of crickets sub­sisting on allochthonous aeolian debris on Anak Krakatau. Philosophical Transactions of the Royal Society of London B 322, 481–486.
83. Thornton, I.W.B. and New, T.R. (1988). Freshwater communities of the Krakatau islands. Philosophical Transactions of the Royal Society of London B 322, 487–492.
84. Thornton, I.W.B. and New, T.R. (1988). Krakatau invertebrates: the 1980s fauna in the context of a century of colonization. Philosophical Transactions of the Royal Society of London B 322, 493–522.
85. New, T.R. and Thornton, I.W.B. (1988). Epipsocetae (Psocoptera) from Peru. Studies on the Neotropical Fauna and Environment 23(4), 225–250.
86. Cole, P.J., New, T.R. and Thornton, I.W.B. (1989). Psocoptera of Flinders, King and Deal Islands, Bass Strait. Journal of the Australian Entomological Society 28, 31–38.
87. Vaughan, P.J., Thornton, I.W.B. and New, T.R. (1989). The Psocoptera of the Krakatau Islands, Indonesia. Treubia 30(1), 1–93.
88. Thornton, I.W.B. (1990). Psocoptera (Insecta) of the island of Moorea, French Polynesia, and comparisons with other Pacific island faunas. Bulletin du Muséum National d’Histoire Naturelle, Paris 4 ser., 11, A(4), 783–828.
89. Endersby, N.M., New, T.R. and Thornton, I.W.B. (1990). Psocoptera from the Grampians and Mt. Arapiles, Western Victoria – a biogeo­graphic analysis. Journal of the Australian Entomological Society 29, 215–224.
90. Thornton, I.W.B., Zann, R.A. and Stephenson, D.G. (1990). Colonisation of the Krakatau islands by land birds and the approach to an equilibrium number of species. Philosophical Transactions of the Royal Society of London B 328, 55–93.
91. Tidemann, C.R., Kitchener, D.J., Zann, R.A. and Thornton, I.W.B. (1990). Recolonization of the Krakatau Islands and adjacent areas of West Java, Indonesia, by bats (Chiroptera) 1883–1986. Philosophical Transactions of the Royal Society of London B 328, 121–130.
92. Thornton, I.W.B., New, T.R., Zann, R.A. and Rawlinson, P.A. (1990). Colonization of the Krakatau Islands by animals: a perspective from the 1980s. Philosophical Transactions of the Royal Society of London B 328, 131–165.
93. Thornton, I.W.B. (1990). Psocoptera of the Hawaiian Islands. Part IV. The endemic genus Palistreptus (Elipsocidae): systematics, distribution and evolution. Bishop Museum Bulletin of Entomology 4, 1–57.
94. Smithers, C.N. and Thornton, I.W.B. (1990). Systematics and distribution of the Mela­nesian Psocidae (Psocoptera). Invertebrate Taxonomy 3, 431–468.
95. Vaughan, P.J., Thornton, I.W.B. and New, T.R. (1991). Psocoptera from Southern Sumatra and West Java, Indonesia: source faunas for colonization of the Krakatau Islands. Treubia 30(2), 103–164.
96. Thornton, I.W.B. (1991). Replacement name for Aaroniella badonelli Thornton (Psoco­ptera, Philotarsidae). Bulletin du Muséum National d’Histoire Naturelle, Paris 4 ser.13, A (3–4), 483.
97. New, T.R. and Thornton, I.W.B. (1992). The butterflies (Insecta, Lepidoptera) of Anak Krakatau, Indonesia: faunal development in early succession. Journal of the Lepido­pterists’ Society 46(2), 83–96.
98. Thornton, I.W.B. and Browning, J.A. (1992). Myopsocidae (Insecta: Psocoptera) from Java, including a discussion of the known Indonesian species. Revue Suisse de Zoologie 99(2), 343–367.
99. New, T.R. and Thornton, I.W.B. (1992). Colonization of the Krakatau Islands by invertebrates. GeoJournal 28(2), 219–224.
100. Rawlinson, P.A., Zann, R.A., Van Balen, S. and Thornton, I.W.B. (1992). Colonization of the Krakatau Islands by vertebrates. Geo­Journal 28(2), 225–231.
101. Thornton, I.W.B. and Walsh, D. (1992). Photographic evidence of rate of develop­ment of plant cover on the emergent island Anak Krakatau from 1971 to 1991 and impli­cations for the effect of volcanism. Geo­Journal 28(2), 249–259.
102. Thornton, I.W.B., Ward, S.A., Zann, R.A. and New, T.R. (1992). Anak Krakatau – a coloni­zation model within a colonization model? GeoJournal 28(2), 271–286.
103. Thornton, I.W.B. (1992). K.W. Dammerman – fore-runner of island equilibrium theory? Global Ecology and Biogeography Letters 2, 145–148.
104. Endang, S.K. and Thornton, I.W.B. (1992). Psocidae (Insecta: Psocoptera) from the islands of Bali and Lombok, Indonesia. Treubia 30(3), 319–379.
105. Thornton, I.W.B., Ward, S.A., Zann, R.A. and New, T.R. (1993). The Anak Krakatau Ques­tion. GeoJournal 29(4), 421–425.
106. Thornton, I.W.B., Zann, R.A. and Van Balen, S. (1993). Colonization of Rakata (Krakatau Is) by non-migrant land birds from 1883–1992 and implications for the value of island equilibrium theory. Journal of Bio­geography 20, 441–452.
107. Maeto, K. and Thornton, I.W.B. (1993). A preliminary appraisal of the braconid (Hymenoptera) fauna of the Krakatau Islands (Indonesia) in 1984–1986, with comments on the colonizing abilities of parasitic modes. Japanese Journal of Entomology 61(4), 787–801.
108. Schmidt, E.R. and Thornton, I.W.B. (1993). The Psocoptera (Insecta) of Wilsons Promontory National Park, Victoria, Aus­tralia. Memoirs of the Museum of Victoria 53(2) (1992), 137–220.
109. Schmidt, E.R., Thornton, I.W.B. and Hancock, D. (1994). Tropical fruitflies (Diptera: Tephritidae) of the Krakatau Archi­pelago in 1990 and comments on faunistic changes since 1982. Ecological Research 9, 317–324.
110. Compton, S.G., Ross, S.J. and Thornton, I.W.B. (1994). Pollinator limitation of fig tree reproduction on the island of Anak Krakatau (Indonesia). Biotropica 26(2), 180–186.
111. Thornton, I.W.B., Ward, S.A., Zann, R.A. and New, T.R. (1994). Further comments on the Anak Krakatau Question. GeoJournal 33(4), 493.
112. Thornton, I.W.B. (1994). Figs, frugivores and falcons: an aspect of the assembly of mixed tropical forest on the emergent volcanic island, Anak Krakatau. South Australian Geo­graphical Journal 93, 3–21. (Based in part on the Brock Memorial Lecture, given to the Royal Geographical Society of Australasia [South Australia Branch] in August 1993.)
113. Wang, Q., Thornton, I.W.B. and New, T.R. (1994). Systematics and biogeography of the Australian-New Guinean genus Thoris Pascoe (Coleoptera: Cerambycidae: Phora­canthini). Invertebrate Taxonomy 8, 839–860.
114. Thornton, I.W.B., Partomihardjo, T. and Yukawa, J. (1994). Observations on the effects, up to July 1993, of the current erup­tive episode of Anak Krakatau. Global Ecol­ogy and Biogeography Letters 4, 88–94.
115. Schedvin, N.S., Cook, S. and Thornton, I.W.B. (1994). The diversity of bats on the Krakatau Islands in the early 1990s. Bio­diversity Letters 2, 87–92.
116. Wang, Q., New, T.R. and Thornton, I.W.B. (1995). Phylogeny and distribution of the phoracanthine genus Atesta (Coleoptera: Cerambycidae) from Australia. Systematic Entomology 20, 229–238.
117. Wang, Q., Thornton, I.W.B. and New, T.R. (1996). Biogeography of the phoracanthine beetles (Coleoptera: Cerambycidae). Journal of Biogeography 23, 75–94.
118. Thornton, I.W.B., Compton, S.G. and Wilson, C.N. (1996). The role of animals in the colo­nization of the Krakatau Islands by Ficus species. Journal of Biogeography 23, 577–592.
119. Ward, S.A. and Thornton, I.W.B. (1999). Guest Editorial. Equilibrium theory and alternative stable equilibria. Journal of Bio­geography 25, 615–622.
120. Runciman, D., Cook, S., Riley, J., Wardill, J. and Thornton, I.W.B. (1999). The avifauna of Sebesi, a possible stepping-stone to the Krakatau Islands. Tropical Biodiversity 5(2), 1–9.
121. Wang, Q., Thornton, I.W.B. and New, T.R. (1999). A cladistic analysis of the phora­canthine genus Phoracantha Newman (Coleoptera: Cerambycidae: Cerambycinae), with discussion of biogeographic distribution and pest status. Annals of the Entomological Society of America 92(5), 631–638.
122. Smithers, C.N., Peters, J.V. and Thornton, I.W.B. (2000). The Psocoptera (Insecta) of Norfolk and Philip Islands: occurrence, status and zoogeography. Pro­ceedings of the Linnean Society of New South Wales 121, 101–111.
123. Ward, S.A. and Thornton, I.W.B. (2000). Chance and determinism in the development of isolated communities. Global Ecology and Biogeography 9, 7–18.
124. Thornton, I.W.B., Mawdsley, N.A. and Partomihardjo, T. (2000). Persistence of biota on Anak Krakatau after a three-year period of volcanic activity. Tropical Biodiversity 7, 25–43.
125. Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. I. General introduction. Jour­nal of Biogeography 28, 1299–1310.
126. Harrison, R.D., Banka, R., Thornton, I.W.B., Shanahan, M. and Yamuna, R. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. II. The vascular flora. Journal of Biogeography 28, 1311–1338.
127. Schipper, C., Shanahan, M., Cook, S, and Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. III. Colonization by birds. Journal of Biogeography 28, 1339–1352.
128. Cook, S., Singadan, R. and Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. IV. Colonization by non-avian vertebrates. Journal of Biogeography 28, 1353–1364.
129. Shanahan, M., Harrison, R.D., Yamuna, R.Y., Boen, W. and Thornton, I.W.B. (2001). Colo­nization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. V. Colonization by figs (Ficus spp.), their dispersers and pollina­tors. Journal of Biogeography 28, 1365–1378.
130. Edwards, J.S. and Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. VI. The pioneer arthropod community of Motmot. Journal of Biogeography 28, 1379–1388.
131. Thornton, I.W.B., Cook, S., Edwards, J.S., Harrison, R.D., Schipper, C., Shanahan, M, Singadan, R. and Yamuna, R. (2001). Coloni­zation of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. VII. Overview and discussion. Journal of Biogeography 28, 1389–1408.
132. Endang, S.K., New, T.R. and Thornton, I.W.B. (2002). The Psocidae (Psocoptera) of Java and the eastern islands of Indonesia. Inverte­brate Systematics 16, 200–276.
133. Thornton, I.W.B., Runciman, D., Cook, S., Lumsden, L., Partomihardjo, T., Schedvin, N., Yukawa, J. and Ward, S.A. (2002). How important were stepping stones in the coloni­sation of Krakatau? Biological Journal of the Linnean Society 77, 275–317.

Miscellaneous publications

1. Thornton, I.W.B. (1983). J. Linsey Gressitt (1914)–(1982). GeoJournal 7(6), 481–482.
2. Thornton, I.W.B. (Ed.) (1985). 1984 Zoolog­ical Expedition to the Krakataus. Preliminary Report. La Trobe University Department of Zoology. Miscellaneous Series No. 1, 57 pp.
3. Thornton, I.W.B. (Ed.) (1986). 1985 Zoo­logical Expedition to the Krakataus. Prelimi­nary Report. La Trobe University Department of Zoology. Miscellaneous Series No. 2, 63 pp.
4. Thornton, I.W.B. (1986). Krakatau Rebirth. Australian Geographic 1(2), 40–54.
5. Thornton, I.W.B. (1987). A Guide to Kraka­tau. Sponsored by the Krakatau Foundation and P.H.P.A. (Forest Protection and Nature Conservation, Indonesia). 21 pp.
6. Thornton, I.W.B. (Ed.) (1987). 1986 Zoolog­ical Expedition to the Krakataus. Preliminary Report. La Trobe University Department of Zoology. Miscellaneous Series No. 3, 59 pp.
7. Thornton, I.W.B. (1989). The recolonisation of Krakatoa by animals. Pacific Science Asso­ciation Information Bulletin 41(3), 13–23.
8. Thornton, I.W.B. (1990). Message from Melbourne. Bio News 28, 4–6.

Ian Ritchie AO BA MA MEng PhD ScD FTSE FRACI FAIMM; scientist, engineer, teacher and humanist, brought fresh understanding and relevance to the relationships between metals and fluids through his work on metal oxidation, electrochemistry and hydrometallurgy. 

Passionate about education, society and the environment, he constantly sought new ways to interest young people in science and its role in the future wellbeing of human society and the fragile Earth. He served the community with energy and dedication as a valued advisor to government, industry and academia. 

He contributed much to the establishment of air quality standards in Western Australia, and is credited with preventing the dismantling of the WA Government Chemical Laboratories and steering them to rebirth as the ChemCentre. 

He was the founding CEO of the AJ Parker CRC for Hydrometallurgy, which became the world's foremost centre for hydrometallurgical research under his inspirational leadership. 

Ian was a kind and humble man with huge talent who was always ready to share his wisdom. He was a passionate and prolific scientist, but his greatest joy came from his family – Ann, the love of his life, and his three wonderful children, Kathy, Andrew and Alex. 

Driven by a deep sense of fairness, he railed against injustice and stupidity wherever he saw it. All who knew him will miss his ready wit, awesome erudition and endless creativity, but the imprint of his contribution and influence will never fade. He was inducted to the Western Australian Science Hall of Fame in 2016.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 28(1), 2017. It was written by Gregory P. Power, M. Ann Ritchie, Kitty J. Drok and Ian D. Macleod FTSE.