Professor Helene Marsh was awarded a BSc Hons from the University of Queensland in 1968 and a PhD from James Cook University of North Queensland in 1973. In 1973 she began her lifelong work on marine mammals and their habitat, focusing initially on dugongs. As well as studying marine and coastal animals she also started a longitudinal study of Black Rock wallabies in 1986. This study is still ongoing. Professor Marsh was awarded a Personal Chair in Zoology at James Cook University of North Queensland in 1991 and became Professor of Environmental Science at this same institution in 1994. She served as the chair of the Great Barrier Reef Consultative Committee from 1998 to 2000. In 1998 she was the recipient of an international Pew Charitable Trust Award for marine conservation. Professor Marsh is currently leading a program at the Cooperative Research Centre for the Great Barrier Reef World Heritage Area (CRC Reef Research Centre) that is looking for sustainable solutions to human impacts on the Great Barrier Reef.

Interviewed by Dr Hugh Tyndale-Biscoe in 2000.

Contents

• Family support
• An attraction to biology and research
• Life-changing developments
• Marine elephants, manatees and terrestrial dugongs
• The sustainability of traditional dugong catches
• By-catch threats to marine wildlife
• 'Think about losing 1000 square kilometres of habitat'
• 'Marine koalas': weighing lifestyles and conservation
• The can of worms: bio-political connections
• Synergies between dugong and manatee research
• A move into cetacean research
• Amazing implications of whale age–sex research
• Some consequences of producing unwelcome data
• Rock wallabies: the mammal research moves inland
• Is there an ideal blend of research skills, curiosity and creativity?
• Peer review, integrity and challenges to responsible research publication
• Women scientists' balancing act

Family support

Helene, it's a great pleasure to interview you for the Academy series. First of all, what is your family background? Were you supported much by your parents?

Yes, very much. My parents were both well educated. My mother trained as a teacher before the war and then went into the army during the war as an education officer, in fact the only woman army education officer in the Northern Territory. (I think she had a very independent view about what women could do, although she probably wouldn't like to be called a feminist.) And later my mother gained a masters degree. My father had degrees in law and economics, probably quite an unusual combination at the time. As a child I definitely was encouraged to do well academically, and for as long as I can remember it was always expected that we would go to university. That was never in doubt – learning, knowledge, was just incredibly valued in my family. One of my brothers is a professor of English at the University of London. My other brother makes films and videos; he was always considered the odd one out because he didn't excel academically, but he is very creative.

There was no question that you would get as much opportunity as your brothers?

Well, my father died when I was 13, so I don't know what his views were. But I think my mother expected that I would have an excellent education, go to university, and probably get married. She might have found it surprising for me to have the sort of career I have had. I think that for a long time, particularly when I had young children, she had mixed feelings about that, but if so she seems now to have forgotten those feelings. I think she believed that women should have excellent education but perhaps not the same sort of career commitment as, say, my brothers. In fact, for a long time when I was at school I wanted to do medicine, and my mother used to say, 'Oh, it's such a long course for a girl.' That now seems a ridiculous thing to say, but it did influence me at the time.

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An attraction to biology and research

What got you started as a biologist? Was it family influence, or teachers at school?

Neither, really. I went to the University of Queensland with the aim of doing a science degree with a joint major in psychology and physiology, and as a prerequisite for physiology I did first year zoology. By the end of the first year I decided that psychology definitely wasn't for me but I was interested in biology. I was particularly inspired by the field excursion to Moreton Bay that we undertook that year. In second year I thought I would continue with biology with a view to doing genetics, but again I was rather seduced by the field work. And so by the end of my third year I wanted to do honours in a more field related project – on cone shell venoms, actually, and quite different from what I've done since then.

Did your honours supervisor provide a role model as a research biologist?

My supervisor was Dr Bob Endean. I liked the combination of the field work and the lab work – and maybe I still do – and the possibilities for practical applications of the work he was doing. He was working on venoms. Cone shells, particularly the species that are fish eaters, are very venomous to people, and they have remarkable physiological actions on mammals. I was very attracted to the thought that these venoms might help us understand neurological function, although I don't think that his work did particularly have those insights. (I have noticed in recent years, actually, that some people are using them again in that way.)

And you never doubted that you would have a research career?

Not from the time I was in first year at university, when we had the opportunity to do a small psychology research project. What I did for that was quite creative but quite outrageous when I look back; no first year would ever be allowed to do such a thing now. We had done an incredibly primitive laboratory exercise, looking at skin sensitivity by testing the ability to differentiate between two points when they are touched onto various areas of the body. I had the idea to see whether profoundly blind and profoundly deaf children would have patterns of skin sensitivity differing from those in normal children. I wrote to the Blind School and the Deaf School in Brisbane, got access to these kids, and did the very crude tests – with normal controls. I spent far too long in the second term of my first year doing this project that wasn't really worth very much, and without ethics approval or anything. But I got some interesting results, particularly with the blind children. From memory, the skin sensitivity on their hands was very different from that of sighted children.

I was very intrigued by this exceptionally crude little experiment, and from then on I wanted to be a research scientist. Maybe we should be allowing first year students – with a little more guidance than I had – to undertake research, because I think some students will be very inspired by the experience.

I wasn't really attracted to doing any more psychology, even though I had done very well in it. There seemed to be too much about it that was 'soft'. Later, however, when I did my honours project in biology, the elements that attracted me were the combination of field and lab work, and the potential for the work to make a difference. That is not to say that I'm not a great admirer of and advocate for basic research. I acknowledge that it is incredibly important, but I always liked the idea that there would be an applied element in what I did.

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Life-changing developments

Did you go straight from the honours degree to do a PhD at James Cook University?

No. I had won a postgraduate award to go either to the West Indies to work on a marine invertebrate problem or to a Canadian marine laboratory, so I hadn't applied for a Commonwealth award in Australia. But then Lachlan and I decided to get married and I had to wait a year to get an award in Australia. I went to Townsville because Lachlan had a job there, and for the first year I worked at CSIRO.

So did your private life change the direction of your work towards dugong research?

Not exactly. I continued on the cone shell work for my PhD. After that, though – quite by chance – I started work on dugongs. I still clearly remember Alister Spain telling me at a dinner party about the work on dugongs that he was doing at the university with George Heinsohn, and my mental response, 'Golly, I'd love to do that.' The thing that attracted me even then was the mix of pure science and potential application, the fact that dugongs were traditionally important to indigenous people. But I didn't say anything to him at that time.

One afternoon in 1973, while I was tutoring in Alister's undergraduate statistics class, he came in and said, 'I've just had an interview with the Australian Research Grants Scheme people' (ARGS became the Australian Research Council, ARC) 'who say that these dugong carcasses that we are getting from the shark nets are incredibly valuable, and we should make sure that we get the maximum value out of every one.' He went on to say that the interesting question as to whether or not dugongs are ruminants could probably be solved by fairly anatomical and histological examination of the carcasses. 'Helene, there's $1000. Would you like to do it?' There I was, a young scientist with a PhD and commitments to a young family – I had one young child and was expecting my second – and even then $1000 wasn't very much money. But I said yes, and I guess that really changed my life: from just this small project I got more and more involved.

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Marine elephants, manatees and terrestrial dugongs

Could you tell us a bit about the biology of the dugong, and about your research?

A dugong is a sea cow, a member of the order Sirenia. Their closest living relatives in the marine sense are the manatees, but in other groups of extant mammals they are the elephants. Dugongs are like marine elephants, I guess.

The Sirenia species are vegetarian. Four species exist now – three species of manatee and the dugong. The manatees are much less specialist herbivores than the dugong. For example, the Florida manatee eats both freshwater and marine vegetation, and about 70 different species of plant. Dugongs, on the other hand, are seagrass specialists and I guess that makes them, like other specialist species, a lot more vulnerable to human-induced disturbance.

Do dugongs occur only in Australia?

No, they have a huge range: they occur from east Africa across to the Solomon Islands, in the waters of more than 40 countries. But Australia is recognised as their stronghold. Australia and Japan are the only two developed countries with dugongs in their waters. Japan has just a remnant population off the coast of Okinawa. (Fascinatingly, the dugong is currently being used by the Japanese as the symbol for a major dispute about the American military presence in Okinawa.) I think Australia has a pretty big international responsibility for dugong conservation; the reports from all of the other countries are not encouraging.

The research on dugongs started, essentially, as carcass analysis. Dugongs were dying in shark nets set for bather protection; they were being eaten by indigenous people for food. Through George Heinsohn, who set up a program of salvaging carcasses, we got material from quite a lot of animals, and after I'd looked at whether or not they're ruminants (they're not) I used that material to start looking at life history. We developed a method of age determination very similar to what had been used for other marine mammals, looking at growth layers in the tusks, which are like growth layers in a tree. They were remarkably clear and we established the periodicity of the rings through an annual increment method: one dark layer and one light layer were laid down per year.

Is that related to the seasonal pattern of the seagrass growth?

We think so. It certainly coincides with that. Seagrass growth is highly seasonal, even in tropical Australia. We then found that dugongs were long-lived and late breeders. They can live to over 70 years of age (as the maximum life span; we've seen very few animals that have lived as long as that) and they probably don't reproduce for the first time before the age of 10. Actually, I have a student doing some more of this work and I think she has an indication that when they're well fed, such as up in Torres Strait, the pre-reproductive period might be a bit less than in some other animals. But she's getting a fair range.

Then we looked at the reproductives. The dugongs had a zonary placenta like elephants and that left conspicuous placental scars in the uterus, so it was pretty easy to identify parous individuals. They also had female reproductive cycles which were incredibly like those of elephants – in Dick Laws' papers on elephants I could almost cross out 'elephant' and write in 'dugong'. And we showed that on average they were only having a calf every three years. Again it was very variable, and in some populations at some times it was much less frequently. I think it's all tied up with the food supply.

How long do they lactate?

We have limited data on that, but lactating females and their calves have been caught in nets and we have aged some calves attending lactating females at 18 months. That all fits in pretty well with the manatee data, except that manatees can breed younger and faster. But they also can eat freshwater vegetation, and they can consume a good deal more in a day than a dugong can. A dugong, being such a specialist feeder, is often quite restricted in what it can eat in a day.

Have you any comments on the recent idea from Roger Short's group that elephants might have been water-living animals?

I loved that. I found it absolutely fascinating, given the paenungulate links with Sirenia. There has been a suggestion that elephants are terrestrial dugongs, which might indeed be a new way of thinking about it. But probably the more plausible explanation is that they have a common ancestor that lived in marshy country, probably on the interface between land and sea. That fits in with the fossil evidence.

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The sustainability of traditional dugong catches

From what you're saying, the dugong has a life span very similar to humans and it reproduces at about the same age. How do the Torres Strait Islanders and other people around northern Australia who use the dugong regard it? Is it a totemic animal to them?

It is certainly highly valued throughout northern Australia by coastal Aboriginal people and Torres Strait Islanders, and for some groups it's a totemic animal. Some groups also value green turtles very highly, but usually when there is access to both dugongs and green turtles, dugongs are valued more. A lot of that is tied up with manhood, especially since catching a dugong (particularly with a spear, from a bark canoe) is quite a challenging thing to do. Early this century, for a man in the western islands of Torres Strait to be considered eligible to marry he had to have killed a dugong. So it's very important to Aboriginality and manhood.

In places such as the western islands of Torres Strait and the remote areas it's also a really important part of the food supply. I had a student who worked on Mabuiag Island for 18 months, and her census of the number of animals taken indicates that the people are taking enough meat to provide about 300 grams per person per day. Now, I'm sure that not everyone on the island is eating 300 grams of dugong meat per day – some would be exported – but that gives you an idea of how important it is. (As well it might be, considering how little there is to buy in the store on Mabuiag Island.)

I gather the status of the dugongs is not very healthy. In the traditional culture of the Islanders and the coastal Aborigines, was their hunting sustainable?

I think it was, but whether it is still so is another question and probably varies from place to place. Certainly there is evidence from Torres Strait of a dugong hunting culture that lasted thousands of years, so it must have been sustainable. Torres Strait is the most important dugong area in the world, with seagrass supporting habitat of the order of 17,000 square kilometres, which is huge. It may still be sustainable in that area – the jury has to be out. There is probably evidence of local depletion, but some of the most important dugong areas there are not easily accessible by indigenous hunters because they're right out in the middle and a long way from land. The hunting culture has changed, but I don't think there is evidence of really serious depletion in Torres Strait.

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By-catch threats to marine wildlife

And around the Queensland coast?

That's a very different situation. On the Cape York coast there is no evidence of serious depletion in the short term – that is, since the mid-'80s when we started developing aerial survey techniques. But on the urban coast of Queensland, loosely defined as Port Douglas south, there is evidence from a number of sources of quite serious depletion. On the anecdotal evidence, in the historical records the numbers of dugongs are so large it's almost unbelievable. Secondly, our aerial survey evidence indicated a decline between the mid-'80s and the mid-'90s, although recently we've done work which suggests that animals have now moved back into the northern part of that area, probably from the more remote regions further north.

We have just analysed a 40-year data set of dugong by-catch in shark nets set for bather protection. (Such a long data set in biology is quite rare, and we got hold of this one only after a lot of debate, the Department of Primary Industries being fairly reluctant to release it.) There are shark nets set at a number of beaches on the Queensland coast with a view to making the beaches safer to bathers, not by providing a wall to stop sharks from coming to the beach but rather as a fishing device to deplete the local population of sharks. Unfortunately, there is a serious by-catch of marine wildlife such as dugongs and dolphins which get tangled in the nets and drown, and our analysis indicates a significant decline in the dugong catch per unit effort – of the order of 8.7 per cent a year, for 40 years.

The program also has a serious by-catch of small sharks. They are certainly not fatal to humans, but I guess shark attacks are such an emotive issue that it would now be very difficult for the government to stop the program. Advocates have said, 'Well, there has been no fatal attack at a meshed beach since it was introduced.' Other data suggests that the catch per unit effort of major man-eating sharks such as tiger sharks off Townsville, for example, has not changed over the 40-year period, and that the tiger sharks are coming into the region to breed. Because they are not resident in the region, the shark nets may be having no effect at all. But shark meshing on the Queensland and New South Wales coasts is a response to the sort of hysteria and emotive media response we saw last week when two people were killed by white pointer sharks off South Australia.

Whether the number of dugongs caught in shark nets is a reliable index of the dugong population, by the way, is debatable. There is no evidence that the dugongs could learn to avoid the nets, but the human use of the beaches over that 40-year period may have meant that dugongs are less likely to use those areas. We have no data on that. But we do have a data set which indicates a significant depletion in catches over a very long period, and when I combine that with the anecdotal information and the aerial survey data and triangulate, adding in the belief of the indigenous people who use that coast that there has been a serious decline, I think the evidence suggests quite a significant depletion.

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'Think about losing 1000 square kilometres of habitat'

Is the problem that people want to bathe at the beaches where the seagrass grows?

In some areas. The causes of the dugong decline are very complicated and difficult to disaggregate, and probably vary in different regions. In the Cairns region, for example, we have the account by Colin Bertram of his visit to that area. He was a biologist who worked on dugongs in the 1960s, and he says that when he went to the Yarrabah community in 1965, those indigenous people claimed to be catching 200 dugongs a year. But now when we've done aerial surveys in that region we have seen dugongs in such low numbers that we can't make a reliable population estimate. In some regions we certainly find evidence of dugongs being caught in gill nets set by commercial fishermen; in other areas there is evidence of significant habitat loss.

Unfortunately we don't have a 40-year data set for habitat, but we do know the effects of some extreme weather events. For example, in 1992 there were two floods and a cyclone in very quick succession in the Maryborough region. The floods themselves were not remarkable, although one was the fifth highest flood this century, but the fact that there were two of them, three weeks apart, was very unusual. And 1000 square kilometres of seagrass in Hervey Bay was destroyed.

By silt formation?

It was deepwater seagrass – that is, seagrass growing near the limits of its light tolerance in greater than 15 metres – and with a prolonged flood plume it just had to die. A seagrass survey had been done only a few months before, and then it was repeated. Well, dead dugongs were found all along the New South Wales coast, and some live ones; about 100 dugongs were recovered in the few months after that event, and more were found that year on the New South Wales coast than in all other years put together. So something very strange was going on.

Think about losing 1000 square kilometres of habitat. It has come back, but the concern is that we have extreme weather events coupled with bad land-use practices. For example, the sediment going into the Great Barrier Reef lagoon per year has increased by four or five times since European settlement. We have dugongs in both intertidal and subtidal areas. The intertidal seagrass is probably okay, because it's getting light when the tide goes out, but the likelihood is that the depth margin of the subtidal seagrass has shrunk. We have no idea, over those sorts of time scales, what habitat loss there has been. But it's hard to believe, given the Hervey Bay story and similar events, that it hasn't been quite serious.

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'Marine koalas': weighing lifestyles and conservation

So two factors have been affecting the dugong population in the last 40 years: the shark nets and the effects on water quality of changes in agricultural practices?

I think it is more than that, Hugh. There are also gill nets set by commercial fishermen for barramundi, in particular. That fishing industry has grown remarkably in the last 50 years. On the other hand, dugongs are no longer caught commercially for oil, as they were in the early part of the century.

The other thing that has probably been important on a local scale is the changed population distribution of Aborigines and Torres Strait Islanders. Before European settlement they would have been strung out along the coast, but now – as a result of government policy – they're concentrated in communities. As a related example, one day an elder on Palm Island (a big Aboriginal community near Townsville) was telling me that they couldn't hunt turtles any more because the turtles were being caught by the trawlers operating offshore. That's probably true to a certain extent, but as I sat and looked at that community I thought that even to have those 3000 people hunting turtles from bark canoes in one bay would never be sustainable.

When people talk about indigenous hunting they always talk about changes in technology, but the complications include changes in the spatial distribution of people. There are now more Torres Strait Islanders living in mainland Queensland than in Torres Strait. Suppose you come down to Townsville as an economic refugee. You've come from dugong city – 17,000 square kilometres of habitat, thousands of animals – to Cleveland Bay, where there are certainly quite a lot of animals but only in the hundreds. No way could the Torres Strait level of hunting be sustainable there.

So all these young men are unable to meet their manhood requirements and marry?

Well, those customs are breaking down now. But the cultural practices are probably very important. There are very complicated reasons for what has happened to dugongs on the urban coast of Queensland, and they probably vary in different areas.

Have you talked with those people in Cleveland Bay about the findings?

I've certainly talked to them. Whether they believe what we tell them is another matter. There has been considerable resistance from the fishing industry about some of our findings. A few years ago the dugong became a symbol for concerns by the conservation movement about fishing. They demanded action on the aerial survey data that indicated a decline, and there were lots of demonstrations about it. In particular, they wanted commercial gill netting banned from the Great Barrier Reef region. This is a broader agenda than just dugongs, however: many conservationists want all commercial fishing banned from the Barrier Reef region. The Commonwealth Minister, Senator Hill, responded to these concerns with quite strong intervention: he introduced 'dugong protection areas' in which gill netting was banned, and refused to issue permits for dugong hunting by indigenous people south of Cooktown. And 38 fishing businesses were closed. The fishermen were compensated, but for many of these people it's a lifestyle issue and they felt very angry indeed about it.

Have you been involved in any other controversy over development areas?

Yes. Dugongs were seen very much as a symbol for Oyster Point, the Port Hinchinbrook development. I think the debate was really about something else, in that the conservation groups were affronted about the Hamilton Island development, which is a product of the same developer as at Hinchinbrook and is seen as very unsympathetic to the environment. But because people find it unacceptable to debate on the grounds of aesthetics, there was a lot of fuss about the dugongs' seagrass disappearing locally. And dugongs became a symbol for this campaign – in effect, 'marine koalas'.

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The can of worms: bio-political connections

How do you equip your students with political science as well as biology for their dugong research?

It's actually quite difficult, and it depends greatly on the student. For example, some of my students have worked extensively in indigenous communities and probably know more about the protocols required than I do. One grew up in Papua New Guinea of mixed racial background, and she is excellent in an indigenous community. Others I wouldn't let near an indigenous community because they tend to have such a strong pro-conservation stance that they would be quite doctrinaire in their dealings with the people, and that can be very inappropriate.

I talk a lot to my students about the political dimensions of these problems, and some are much more receptive than others. One of my former postdocs, who wears his heart on his sleeve and has a very 'green' view, recently completed a report for the Great Barrier Reef Marine Park Authority. He insisted that a photograph of a boat travelling very close to a dugong – clearly, the captain of the boat was oblivious – had to be on the front of the report. But the Marine Park Authority were advised that if they went ahead and released the report with this photograph on the front, they could be sued by the owner of the boat. There is now an ongoing debate between the Reef Cooperative Research Centre and this person, who is not prepared to compromise in relation to the political reality of placing the photograph like that. I personally think he's unwise: he could make his point in other ways but with fewer legal implications. People do differ in how they react to the political nature of the problem.

Does your School of Tropical Environment Studies and Geography have any law as a course component?

Yes, we have a subject in environmental law and policy, which is taught by someone from the School of Law at James Cook University. So our undergraduate students are exposed very much to the broader dimensions of environmental policy – but not all graduate students, of course, have such a background.

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Synergies between dugong and manatee research

You have attended overseas conferences on manatees. What have you been able to offer from your Australian experience in dugong research?

Thanks to assistance and advice from Graeme Caughley, we developed aerial survey techniques which were probably more sophisticated than those of the manatee researchers. I'm not totally certain that the way we do it is directly applicable to manatees because they have a more linear distribution, using a very narrow area around the coast, and the spatial scales are really different. Some of our ideas, though, have been influential. The life history data approach that we used for dugongs was then copied for manatees; the converse was that we were able to adapt for dugongs the techniques that had been developed for satellite-tracking manatees. So there's been quite a bit of synergy, which has helped both groups.

Are manatees common, or are they also threatened by changes?

Well, the absolute number of Florida manatees, just occurring off the Florida coast – which is much smaller than the northern Australian coast – is considerably smaller than the number of dugongs. However, there's increasing data that despite the extraordinary urbanisation of the Florida coast, numbers have actually increased since the '70s – perhaps because we're dealing with a generalist species rather than a specialist one. Florida manatees have teeth very different from dugong teeth, and their jaw opens almost horizontally. Consequently they can feed anywhere in the water column, whereas in dugongs the jaw opens ventrally and they're obligate bottom feeders. So manatees are a totally different animal, and they have actually been able to take advantage of the huge increase in freshwater aquatic vegetation that has occurred concomitant with the urban development. Some animals just seem to cope very well with urbanisation. To use a possum analogy, manatees are more like brush-tails and dugongs might be like some of the specialist rainforest ringtails.

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A move into cetacean research

Let's now turn to the other marine mammals that you've worked with: the porpoises and whales. How did that cetacean work begin?

Again rather by chance. When I first started working on dugong life history and reproductive biology, I had the opportunity to go to an international workshop in La Jolla, California, on age determination in marine mammals. There I met Toshio Kasuya, who was doing a lot of work on cetacean life history based on the Japanese drive fisheries for small cetaceans. Because Japan is so close to ocean environments, it has access to an extraordinary cetacean fauna and these traditional fisheries for so-called small-type whaling have been operating off the Japanese coast since the 15th century at least. The fishermen would drive schools of dolphins or small whales into bays, seal off the bay and kill all the animals in the school.

So when the Japanese say that whale meat is a traditional meat, this is actually true of the small whales?

There was large whaling there too, as a sort of cottage industry. But no-one could say that Antarctic whaling is traditional. I've been to places such as Taiji, which has operated as a whaling village for hundreds of years. In fact, I had the privilege of going to a town meeting where the community were having great problems in dealing with the international criticism – by so many people – of what they had been doing for such a long time. I guess it's the same sort of reaction as timber workers in Australia have had on discovering that what they've spent their life doing is unacceptable. I have often thought how I would feel if being an academic became unacceptable.

Molecular biologists might be in that position nowadays!

You're right, and it can be hard when people do things generally in good faith, often in ignorance of the broader ramifications of their work.

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Amazing implications of whale age–sex research

When Toshio Kasuyo told me, in La Jolla, that he was working on both the reproduction and age determination, I said I was about to start studying dugong reproductive organs but there was nothing written. 'Can you help me?' I asked. His reply was, 'Helene, how about working with me on pilot whale ovaries? You could 'cut your teeth' on ovaries about which a lot more is known, and then go back to the dugong ovaries.'

This was in about 1978. In 1980 I went on a study leave to Canada, and had the opportunity to go to Japan several times and to work with Toshio on the pilot whale ovaries. But because I was based in Vancouver and he was in Tokyo, he did all the age determination completely independently of my looking at the ovaries – which in the light of what happened was a great strength to the study. These pilot whales lived to up to age 63, but when Toshio and I met and put all the data together, we had such big samples that we could calculate that when the whales stopped breeding at age 35 they had a mean life expectancy of 14 years. We had found a post-reproductive stage. I did the histology on the ovaries and they were like post-menopausal human ovaries: there weren't any follicles left. All this was pretty amazing.

This was the first time it had been recorded in a wild mammal, wasn't it?

That's right. Even more amazingly, some females were still lactating into their early 50s, even though they had post-reproductive ovaries. We don't know whether they were suckling their own calves, but when we analysed the age composition of all the schools it was certainly plausible that they were.

Then Peter Best, in South Africa, who had done work on sperm whales, got permission to take some sperm whale calves under scientific permit. Sperm whales are another long-lived group of tooth whales that live in matrifocal kinship groups. That is, the females are very closely related. They stay in the same group for life, with their mothers, and the males – the sons – move away. (The genetics that has subsequently been done on pilot whales, by the way, substantiates that.)

Apparently, when sperm whales were being hunted for commercial whaling there was a debate about how long their mothers suckled their young. But because even quite young sperm whales eat fish and squid, just looking at what is in their stomachs cannot indicate whether they're still suckling. So Peter tested for lactose. He found that male sperm whales were suckling up to age 13 and females up to age nine. Whether the males are suckling for that very long time from their mothers or whether there is communal suckling isn't definitely known, but when we put that together with the pilot whale data and then looked again at the school analysis to correlate the age–sex composition of these schools with these very old lactating females, the most plausible explanation was that they were lactating male calves up to, say, age 13.

Were the male pilot whales living as long as the females?

No. That's something that has always intrigued me. The differences between human male and female life spans are always attributed to modern stresses on the male, but when you actually review it across mammals – and there are species that are sexually dimorphic – it's usual for the larger sex not to live as long as the other sex. We aged the oldest of the male whales at about 45, and once they reached sexual maturity they had a really different mortality profile than the females. They reached sexual maturity much later than the females: not until the late teens, whereas the female pilot whales were reaching sexual maturity about nine. It seemed the mothers or other females were feeding milk to the male whales right up until they reached sexual maturity, and then they were several times bigger than the females.

We don't know the nutritional advantage of this long lactation, but being a pilot whale could be quite complicated, with a lot of learning involved. There is probably a link between this very long association with the mother and a very long learning period. There seemed to be a trade-off for female pilot whales between bearing and rearing. Only the older females – particularly the ones who had reached their last offspring – were suckling for such long periods, investing a huge amount in each offspring. The young females were calving much more frequently, probably investing much less. And data from human populations suggests a similar pattern.

Female elephant seals do the same sort of thing, supporting a male offspring for much longer because it's got a much longer growth period.

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Some consequences of producing unwelcome data

Has the political debate about whaling impinged on you as a result of this work?

Not directly. But my colleague, Toshio Kasuya, no longer has access to this material, because the fishermen in Japan don't like him and so he now works for a university, not the Fisheries Agency. He still goes to International Whaling Commission meetings with funding from the Whaling Commission themselves, but it is really difficult for him to go because the Japanese government always object. They believe that his data don't support the government stance on whaling. He and I collaborate on dugong work (in which he has been involved in Japan) but not on pilot whales any more. And I haven't been directly involved with the whaling debate.

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Rock wallabies: the mammal research moves inland

I understand that when you became Professor of Environmental Science you wanted a study site on land, and that was the beginning of the Black Rock wallaby study.

Yes. This was really at Rhondda Jones's instigation. Marine mammal work is logistically difficult and expensive for graduate students, and she was very keen that I set up a terrestrial project. Also, after doing the pilot whale work I was quite intrigued by the idea of setting up a longitudinal study, tracing individuals through time. Black Rock is an isolated outcrop which supports quite a large population (it varies between about 60 and 100) of rock wallabies, Petrogale assimilis. If you're going to conduct a long-term study like that it is less controversial to work on a relatively common species like Petrogale assimilis than on an endangered species.

The Black Rock site is many kilometres from the nearest such outcrops, and we would have thought that the population was very isolated. But the genetics done by one of my students, Peter Spencer, shows that the colony has very high genetic diversity, so there must be far more connectivity with the other colonies than we've ever been able to establish directly. Another of my students did a lot of radio tracking work at the site and never recorded feeding rock wallabies going more than 700 metres from the rock, yet the genetics don't support that pattern at all. So there's a really interesting question about dispersal from Black Rock. The study started in 1986 and is still going, but just in maintenance mode now: we're checking it a couple of times a year. I would like another student to work on that dispersal problem, particularly, although it's a very hard problem to crack.

The fourth PhD from Black Rock is very close to completion and will be the most exciting. Steve Delean, who is statistically very competent indeed, has taken the longitudinal data set and looked at the patterns over time. (Steve worked as a research officer on the project for a while, so he collected some of the data himself, but he has also used the long-term data collected by everyone else.) He has shown that the size of the population is almost totally explained by variations in the climate. When we have good years, the population booms; when there is a series of droughts, there is almost no recruitment into the colony.

But also Steve has found something absolutely fascinating about the survivorship of the pouch young, before pouch emergence. Correcting back for other components like the age of the mother, the season and the climate at the time of conception – these are not seasonal breeders – he has found very strong differential survivorship of pouch young of males and females, depending on the season and the climate at the time. When the going is very tough, the male pouch young have a much lower survivorship than the females; when the going is very good, the situation is reversed and male survivorship improves incredibly. That fits in exactly with what one would expect on theoretical grounds, and also with the sorts of things that Clutton-Brock found for red deer. But the nice thing about the rock wallabies is that their long period of pouch life, in which they're accessible, has allowed Steve to see exactly when this differential survivorship occurs. It's during the second stage, which is very interesting.

The study has been going almost for the full life of a wallaby, which is normally about 15 years. Are you going to continue it?

It's getting much harder, which is why I really would like to have another student working there. John Winter has become involved in the study and we have applied for money through the Savanna Cooperative Research Centre, so we hope it's re-funded. They're interested in long-term sites and it's very hard to keep long-term studies going, but that's a pretty valuable one. The overwhelming influence of climate in that environment is probably not surprising, but it is good to actually have the data.

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Is there an ideal blend of research skills, curiosity and creativity?

Helene, after being appointed to a personal chair for research and being involved with students in these fascinating studies, you're now getting more involved in top management. How are you finding the change to university manager?

It's a bit of a problem. I probably enjoy everything I do, which makes choices harder. I very much enjoyed being head of the School of Tropical Environment Studies and Geography, but I doubt that I will ever want to be head of school again. I don't miss fighting with the dean over money, for example. Being dean of postgraduate studies for the university is about policy rather than line management. It is really fascinating during these very turbulent times for research training in Australia, with very dramatic government intervention. Some of it is very good; some is awful.

The good aspect is that the funding for research training is being made much more transparent and universities are being required to respond by spending that money on research training rather than on underwriting undergraduate work as in these recent tight years. We're also being asked to ensure that supervisors are trained. That's good, because while some people are excellent supervisors without training, others are not. I suspect that some people are beyond improvement in teaching, but there is certainly potential to make a lot of people a lot better.

We are also being asked to look at skills development for research students, largely because many them will not become academics or researchers but will use their postgraduate training in other ways. Also, because everybody is getting busier and busier, the one-to-one apprenticeship model which we've always favoured in this country may not be working as well as it did when time was at less of a premium. So we must provide students, in a rather more organised way, with the opportunity to get the training and the skills they need for both their project and their career. When students come in, we really have to look at their needs. For example, if they're going to need multivariate statistics to analyse their data in year three, we have to do something to make sure that they have those skills in advance. I guess there's enough of a missionary in me to be quite attracted to the challenge of implementing excellence in research training. Because I have had a lot of students myself, I'm very committed to excellent research training.

To an older generation, training seems almost the antithesis of research. I think that in your own career the ideas and the questions have come first and then you've got the tools to solve them – just as in the traditional view of research. Does the increased emphasis on necessary training diminish the curiosity side of research?

I think it does, and that is a major concern. I worry to see more and more research scholarships tied to specific projects. Like you, I routinely review for the Australian Research Council. I get concerned when an ARC project is developed in enormous detail and we are told that most of the work will be done by a student who will start on 30 January, say, and go out collecting data as from 5 February. (I have actually seen such a thing in a proposal.) Where does that leave the student? Is the student being turned into a research assistant? I don't like that. I am not opposed to the idea of some of our students doing projects where the topic is predetermined, but I hate it when the project is determined in incredible detail. Students have to put their own stamp on the work. We're training creative researchers, not automatons.

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Peer review, integrity and challenges to responsible research publication

Could we turn from influences on research training to influences on the way the research results are published. With funds coming now from outside stakeholders, there may be a tendency to think it is necessary only to write the report for the stakeholder, not to publish in refereed journals. Is peer review still of value?

Peer review is absolutely essential. Even when I have to write reports, I now insist on peer review as part of the contract. Some government agencies require peer review to protect them; I want peer review to protect me and my integrity as a researcher. When my research has led to controversial policy, I have found that the people impacted by the policy have reacted by hiring their own researchers with the specific agenda of destroying the research reputation and destroying the research on which the policy is built. I feel very, very strongly that the only protection against that is peer review.

I believe that when peer review standards are paramount, that fact colours the whole research process right from the beginning: you design all the research with the aim that it will satisfy an impartial judge. Is that your view?

I agree. I support Cooperative Research Centres, but I am seeing the flip side. For example, the government agency most involved with the Reef CRC, the Marine Park Authority, see research as providing the information they want, often for political ends. I think that an agency which asks for the research, the information, that it wants will be less concerned about the process of research and the rigour of the process. When you know something is going to be subjected to peer review, your integrity as a scientist becomes very important and you're not going to compromise the process. But I feel we are under increased pressure to compromise process.

I am concerned also that a set of data can be interpreted in different ways depending on the theoretical paradigm of the researcher. (This is acknowledged in the code of conduct of the Ecological Society of Australia.) I've certainly found that fisheries scientists and conservation biologists interpret data quite differently. Regardless of who might be right or wrong there, we have to acknowledge that those different paradigms exist and that research presented only as information can very much be misinterpreted. We do need to have those debates and it is important that different people might look at the same set of data in different ways. I am concerned that when research is seen purely as information we'll lose this. It is too easy to select information that supports your argument – or, more scarily, to bury the information that doesn't support your argument.

It is really important to make young scientists more aware of the dangers of their research being used wrongly. I think they often have too idealistic a view of research. I certainly found, when my research was criticised in a pretty nasty way by the fishing industry, that I was very unprepared for such an attack. But senior colleagues who had been involved in the forest debate were surprised at my naivety. We don't talk nearly enough about these broader dimensions of research, but unless we do, other researchers will be burned. So I would like us to be talking about these problems.

I have a Pew Fellowship in marine conservation. The Fellows get together every year at wonderful meetings of the Pew Foundation to talk about their work and its broader dimensions, but even in that forum I find that scientists who have never been exposed to the darker side of science tend to be very dismissive of the concerns of those that have. One of the Pew Fellows is a woman who works on Pfiesteria, a toxic marine dinoflagellate about which there have been programs on Australian television. There was evidence that the effluent from pig farming in Chesapeake Bay in the United States was influencing the outbreaks of Pfiesteria, but of course the pig farmers didn't want to know about it: the researcher was subjected to the most extraordinary abuse. (She has since been vindicated.) We have to prepare young scientists to recognise that these things can happen.

There is a long tradition of challenging science with calumny. Rachel Carson was pilloried by the chemical industry when she wrote her original book, Silent Spring. Do you address this problem in your postgraduate courses?

I do, actually. I give examples and I talk about it. I also talk to students a lot about the continuum from being what I call an analyst to being an advocate. I tell them that where they want to be on that continuum is up to them, but I certainly warn them about taking the extreme advocate position, because I believe that scientists who do that usually diminish their reputation as scientists. I don't tell them not to do it, but I tell them to think very carefully if they're going to do it.

We come back to peer review, don't we?

Absolutely. Peer review is absolutely critical to the standards in science.

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Women scientists' balancing act

Let's look at one more question facing some students. You had a very supportive family who didn't see that being a girl precluded being a research worker. But for others there have been big problems. What is it like for your women students today?

I think women are now less confused about the potential for a dual role. Whereas for a long time I felt quite confused about how I could undertake a dual role, or even whether I should, I think my women students have no doubts. On the other hand, some of them seem not very pragmatic about managing the complexity of that dual role. The fact that I had a child while I was a PhD student seems to have been a licence for a number of my students to do likewise, but many of them have probably done it in a much less supportive environment than mine was. Some of them may have been quite unrealistic about what they could and couldn't do. So the belief that you can do everything does have a flip side. People do need to be a bit pragmatic and look at what's realistic.

This has been a most interesting day, Helene, and I hope that many other people will take heart from what you've been saying. Thank you very much.

Thanks, Hugh. I've enjoyed it.

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Jacques Miller attended St Aloysius' College in Sydney then went on to study medicine at the University of Sydney. During his medical studies, he took a year off in 1953 to do a BSc in a bacteriology laboratory. Miller did his medical residency at Royal Prince Alfred Hospital, Sydney and was then awarded a Gaggin Fellowship to do medical research in London. At the University of London, he completed a PhD in 1960. In 1963 he spent a year at the National Institutes of Health in Maryland. He returned to London for several years and in 1966 moved to Australia to become head of the experimental pathology unit at the Walter and Eliza Hall Institute of Medical research. Miller served on the International Research Agency for Cancer, and was president of the Scientific Council. He served on the World Health Organization in the area of eradicable diseases and had a term on the International Union of Immunological Societies.

Interviewed by Professor Frank Fenner in 1999.

Contents

• An intercontinental childhood
• Into medicine and on to research
• Leukaemia virus and newborn mice
• Immune tolerance: what was the role of the thymus?
• Why did the mice waste away? An unexpected finding
• Spreading the word
• So why did the mice waste away?
• Testing a theory – and a very testing procedure
• Could thymus lymphocytes become immuno-competent?
• Differentiating lymphocyte subsets
• Eventual acceptance, but new complexities
• Studying delayed-type hypersensitivity: could MHC restriction be shown in vivo?
• Immune-responsiveness genes: a masquerade
• Where was that elusive T-cell receptor?
• Why are we tolerant to our own tissues? More complexities
• Contributing to the scientific community

An intercontinental childhood

Jacques, you had an interesting and exciting childhood and early adulthood, moving between France and China before coming to Australia. Could you tell us about that?

My father was the manager of the Franco-Chinese Bank in Shanghai, staying in China and Japan for about 25 years. One of my sisters was born in Paris, before my parents left for Shanghai, another sister was born in Shanghai, and I was conceived in Shanghai but born on holidays in France, where my mother had gone for health reasons. When she was on the boat – that is how people went in those days – she found herself pregnant, so she decided to have the baby in Nice, next to Marseilles, where the boats used to go.

After less than a year we went back to China but about 3 years later my mother returned to France with the family – again for health reasons and because she wanted my elder sister to be educated there, not in China. Unfortunately, my elder sister contracted tuberculosis, and so the family moved to Switzerland for her to be cured. (That was before the days of streptomycin.) My father eventually came to Switzerland for long service leave, but then World War II started and we left in a hurry for Shanghai. After about a year and a half, knowing that the Japanese were going to be involved in the war, we went to Australia. That was about 3 months before Pearl Harbor.

I didn't know much English at all and had to learn very quickly. But children do learn very quickly. Because my father had thought the Jesuits were wonderful in Shanghai, he decided that in Sydney I would go to a Jesuit school, St Aloysius', and my sister went to Loreto Convent. Both were in Milsons Point. Actually, Gus Nossal was a year ahead of me at St Aloysius'. He was always enthusiastic and very outspoken. He loved debating and was very articulate, just like now.

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Into medicine and on to research

You did pretty well at school, I gather.

I was lucky, I did very well at school and therefore I got an Exhibition, which in those days was a kind of fellowship which took you to university. I had always wanted to do medicine. One reason was that having escaped from the war twice, I was dead against violence. I didn't think I could escape being drafted but I thought I might be a doctor in the army rather than a soldier. I was interested in medicine also because when my sister got tuberculosis they didn't know anything about how infectious diseases were conquered by the body – what pathogenetic mechanisms dealt with such infections. That always interested me and I wanted to learn something about it.

In 1956 I did my residency in the Royal Prince Alfred Hospital, which was the main hospital in Sydney. That is where I met my wife, who was a nurse at the time, and we got married the year I was a resident. During my medical studies I took a year off in 1953 to do a Bachelor of Science, because I hoped to do a little bit of research into pathogenetic mechanisms. In medical school you don't learn much about that – you learn the various signs and symptoms but you don't go into depth. I chose to do a BSc (Med) year in Pat de Burgh's bacteriology department, at the University of Sydney, because they were studying the response to ectromelia virus and that sounded very interesting.

An old friend of mine, ectromelia virus, which I worked on with Mac Burnet in the late 1940s. So you gained a taste for research and knew it was what you wanted to do?

Even more than before, yes. After my medical training and internship, I saw an ad in the Medical Journal of Australia about a Gaggin fellowship to be given by the University of Queensland for 2 years' medical research in London and then 1 year in Brisbane. I applied for that fellowship and was very pleased that it enabled me to do a PhD at the University of London – based at the Chester Beatty Research Institute, the institute of cancer research in South Kensington.

When I arrived at the institute I didn't really know exactly what I wanted to do as a PhD subject and I was told to look around. At the main institute in London they were studying chemical carcinogenesis at the chemical and nucleotide levels. I was not terribly interested in this – I was rather depressed, actually. But the institute had satellites, one in Surrey and one in Chalfont St Giles at Pollards Wood, and at Pollards Wood I met R J C Harris, who was interested in the Rous sarcoma virus and its effects. He suggested that I study another tumour virus, such as the leukaemia virus (Gross, in the United States, had just isolated it in tissues) and I decided that the induction of mouse leukaemia by Gross virus would be the subject for my PhD.

So that kept you in virology, but in a cancer research environment.

Yes. Having worked in virology in de Burgh's laboratory, I was very pleased.

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Leukaemia virus and newborn mice

In those days, viruses were not generally accepted as a cause of cancer. Gross had been able to induce leukaemia in mice by using filtered extracts of mouse leukaemic tissues. He had never isolated the virus as such, but the filtered extracts were able to induce leukaemia in low-leukaemic strain mice, that is, strains of mice which normally don't get leukaemia. Leukaemic extracts from high-leukaemic strain mice, which normally do get leukaemia, were injected into newborn mice at birth – it had to be at birth and not later – and the leukaemia developed 3 or 4 months later.

Rather than develop my own virus from the mouse leukaemic extracts, I wrote to Gross saying, 'Would you mind sending me some of your filtered extracts? I have to do a PhD, I've only got 2 years and it would take me too long to start from scratch, whereas if you sent me your filtered extracts I could induce leukaemia and then repassage that through a series of mice.' Luckily for me, he was kind enough to do it.

I remember that Ludwik Gross wrote the book Oncogenic Viruses.

In those days you had to inject the extracts in newborn mice – if you injected into adult mice nothing happened. That was interesting in itself. Knowing that people like Sir Peter Medawar had discovered immunological tolerance by injecting cells into newborn mice, I wondered whether that had anything to do with the induction of leukaemia by injecting these kinds of extracts to newborn mice, and I was very interested in the phenomenon of immunological tolerance. I had not met Medawar but I decided to see if I could learn something from him.

In 1960 Medawar gave the Tercentenary Lecture of the Royal Society. Afterwards, in a quick chat, I asked whether he would be kind enough to teach me a few things about how to induce tolerance in mice. He said, 'Come along one day,' and very nicely he arranged for one of his co-workers, Leslie Brent, to teach me how to inject mice intravenously, how to skin-graft, and other techniques.

Brent later became Professor of Microbiology at St Mary's, didn't he?

That's correct. I was then able to do some experiments on immune tolerance, particularly to see whether I could induce leukaemia in low-leukaemic strain mice simply by putting in them a thymus graft from a high-leukaemic strain. That was a naive idea, but all these techniques that I learned at the time became very useful later.

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Immune tolerance: what was the role of the thymus?

How did you get onto thinking about the thymus?

Lymphocytic leukaemia induced by Gross virus began in the thymus and then spread elsewhere. Many other types of lymphocytic leukaemia in mice do exactly the same thing: you can induce leukaemia in low-leukaemic strains by various agents, such as irradiation or chemical carcinogens like dimethylbenzanthracene. All these lymphocytic leukaemias begin in the thymus and then spread. In fact, the spontaneous leukaemia of high-leukaemic strain mice, which are called AK, also begins in the thymus. In America and other places it had already been shown that thymectomy (removal of the thymus) in older mice would prevent both spontaneous leukaemia and leukaemia that had been induced by the various agents I mentioned, provided it was done at about 1 or 2 months of age – so preventing leukaemia from starting at 9 months of age, as it normally would.

That had not yet been done for the virus-induced leukaemia, so one of the first things I did was to see if thymectomy prevented virus-induced leukaemia. I injected the virus at birth and then thymectomised the mice when they were a month old, and it worked – it prevented leukaemia. The next question was obvious: put the thymus back with a graft and see if leukaemia begins again in the thymus. And it did. That was to be expected, but what was not expected was that when I put the thymus back 6 months after adult thymectomy, thinking that by then surely leukaemia could not start again, it did. That suggested that the virus must have remained latent for all that time.

The next experiment was simply to thymectomise adult mice that had been inoculated at birth with the virus, wait for 6 months – not give them a thymus graft – take their tissues, make an extract of their normal tissues and see if that induced leukaemia. And it did. So that was the logical outcome of the experiment.

If it showed that the virus was generalised throughout the body, not localised to the thymus, was the thymus the critical place where it multiplied, presumably because the lymphocytes there were the ideal cells in which to replicate and produce leukaemia?

Probably that is totally incorrect but it is exactly what I thought. You would have to give the virus at birth because the thymus at that particular stage was producing very large lymphocytes – people did not know then why it produced lymphocytes – and perhaps those lymphocytes were the ideal medium for virus replication. So if you inject the virus at birth, it goes to the thymus, replicates and then migrates to the rest of the body. My idea was that if you thymectomised first, at birth, and then injected the virus immediately afterward, leukaemia would not occur – that was obvious – but when you gave the thymus back later on, as I had done, leukaemia should still not occur because the virus would not have had a chance to multiply.

I can see that. That's what my experiment would have been.

So I had to learn the technique of neonatal thymectomy. I could do it in the adult but it is a bit different in the newborn. Finally I worked it out, and actually it's quite easy.

I remember meeting you at Pollards Wood in 1962 and learning from you how to do a neonatal thymectomy. I never used it in my subsequent work, though.

The one problem about neonatal thymectomy is cannibalism by the mothers, so I had to do a lot of mice. I must thank my wife: she helped me very nicely by making sure that the mothers would not eat the babies after thymectomy, or changing them around until she found some that did not eat babies.

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Why did the mice waste away? An unexpected finding

I became worried because although all the mice that survived the operation were perfectly healthy with their mothers and until after weaning, about 4 to 5 weeks of age, they then suddenly wasted and became sick. This was never recorded after adult thymectomy – you can take the thymus of an adult mouse and it lives perfectly well into old age – but after neonatal thymectomy, generally at 6 to 8 weeks depending on the strain, they started wasting away and many of them died. I couldn't understand that.

I did a post mortem, of course, and I found several things. One was lesions in the liver, which looked as if they were infected by a virus – hepatitis virus, for example. But also I found a deficiency of lymphocytes in the lymph nodes and spleen, which was quite remarkable. At that time Jim Gowans and Peter Medawar had both shown that lymphocytes in the spleen, in the lymph nodes and in the recirculation pool – that is the lymph and the blood – were the cells which could initiate immune responses. (They were called immunologically competent cells.) My mice had a deficiency of those lymphocytes, which led me to think they must surely come from the thymus.

Yes, because the thymus is filled with lymphocytes.

Correct, but in those days it was thought that because the thymus had a lot of dying lymphocytes it was a graveyard for them. Not only that, but thymus lymphocytes were not able to induce immune responses in appropriate recipients as the lymphocytes from the spleen and lymph nodes or the recirculating pool were. And also because adult thymectomy did not interfere with immune capacity, people did not believe that the thymus had anything to do with immunity.

Yet, because of the results that I had obtained, I really wondered whether the thymus was the seat of production of those lymphocytes which would eventually become competent. Maybe they were just immature in the thymus – they had to mature and go out, and then become competent. I wrote that idea up in the Lancet in 1961, after being pushed to write it by Sir Alexander Haddow, who was officially my mentor.

It would have been a great help to have somebody like that to push you to write it up and to make sure that you ended up in a journal like the Lancet.

I had to check my idea, and one way was to see whether these mice were totally incompetent. Having learned the technique of skin-grafting from Medawar's group, I gave those mice skin grafts from many different types of strains and even rats, and they accepted the skin grafts. They never rejected them.

The other thing was to see if those mice could be rescued immunologically, to be able to respond as normal mice and reject the skin grafts, by giving lymphocytes – and by giving a thymus graft. Lymphocytes from the circulation rescued them perfectly well, or putting in a thymus graft also rescued them. Putting two and two together, it looked very much as if the thymus was the source of those lymphocytes.

That totally unexpected outcome of investigations of mouse leukaemia must have changed the direction of your life.

Exactly. It changed my life and I actually had to learn immunology.

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Spreading the word

You were not only learning immunology but creating it, to a large extent. Three people brought developments within a decade: Medawar with immunological tolerance, Gowans with the lymphocyte circulation, and you – the youngest – with neonatal thymectomy and the role of the thymus in the production of lymphocytes and therefore immunologically competent cells. When did you first talk in public about it?

There were some meetings at the British Society for Immunology in Oxford and in London, but the first international meeting at which I gave my results was in Perugia, in 1961 – even before the Lancet paper came out. This was a meeting on tumour viruses rather than immunology and actually my written paper is on my work on leukaemia. But what I gave viva voce was on the effects of neonatal thymectomy, because it was so interesting, and the published discussion of my work was on that and hardly at all on leukaemia. People were interested but still very sceptical.

The second international forum was in February 1962, in New York. I had applied to go to the New York Academy of Science and had been accepted, so I gave very detailed results on neonatal thymectomy and the immune response, to an audience mainly of immunologists and transplantation biologists. That made quite a few headlines – even in Time magazine, as a matter of fact!

That's going a long way in America.

In general people accepted the data, but quite clearly some were still very sceptical because of the knowledge that thymus lymphocytes are not able to induce immune responses and that adult thymectomy does not do anything at all. They wondered whether my mice might be so worn down by infections for one reason or another – perhaps they were dirty mice, or something like that – that it would only happen in my case. People in Holland tried to reproduce the work, I remember, but couldn't. That worried me. By that time I had a technician, so I sent her along to that place in Holland where she taught them how to do the thymectomies and of course it worked.

People in America reproduced my work partly but could not cross barriers like the strong H2 barriers. If mice which were thymectomised had skin grafts put on them from mice which were very similar to them in genotype, they accepted those skin grafts, but they would reject skin grafts from mice which were very distant or from other species like rats. Finally it turned out that all those people were not doing complete thymectomies but only partial ones. Once they were able to do complete thymectomies, all the work that I had done was reproduced exactly.

Such a delicate operative technique has to be performed skilfully to succeed. A newborn mouse is a tiny object, and the thymus is a tiny object within that again.

Yes. And it is very interesting that partial thymectomy doesn't have any effect at all. It looks as if the piece of thymus that remains behind has all the necessary equipment to do whatever the whole thymus has to do.

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So why did the mice waste away?

Did you stay on in America?

No. After the February meeting of the New York Academy of Science I returned to London, but I decided I must do a year in a place somewhere in the world which had germ-free mice, because of the problem with infection. My idea was that my mice got infected because they were immuno-incompetent, not for some other reason. They were immuno-incompetent because they were thymectomised at birth, so if I could only do thymectomy in a germ-free situation, I would never get wasting disease but yet I would get evidence of immuno-incompetence, like failure to reject skin grafts. I found that the National Institutes of Health in Bethesda had germ-free facilities; I applied for an Eleanor Roosevelt Fellowship in 1963 and got it; and I was able to do the work in a germ-free tank at NIH. I showed there that these mice lived on perfectly healthily but yet could not reject skin grafts.

So wasting disease essentially had been due to their lack of control of infection?

That's right.

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Testing a theory – and a very testing procedure

NIH is a wonderful place to work in and your work in the germ-free unit must have been a good experience for you.

It was very good. Also, I did some work with Lloyd Law, who was on the same floor as that unit, and showed that neonatally thymectomised mice were much more prone to cancer-inducing agents than normal mice. I had done that before with chemical agents, but we used the polyoma virus and showed it to be true. Burnet had a theory of immuno-surveillance, that the immune system was essential to survey the body against mutations which occur, for example, in cancer and to get rid of cancer cells. So it was very good to show that if you were immuno-incompetent your threshold for developing carcinomas or different types of neoplasm was very much reduced.

It had been on the right track but didn't have the experimental evidence?

That's right, so it was useful to be able to do all these experiments in NIH. After the Eleanor Roosevelt Fellowship I went back to London in 1964, with a few visits to America – to Denver, for example, where they wanted to thymectomise opossums in the pouch! In those days people only thought of one type of lymphocyte, and the very fact that neonatal thymectomy in mice was still associated with some lymphocyte production suggested that perhaps you should take the thymus out before birth, to make sure that all the lymphocytes have been removed. That is impossible in mice, but in opossums you could do it in the pouch. They had opossums in Denver and were working on them, so I went there twice to do it. It was very difficult and I never got very far. They were like balloons filled with nothing but blood.

It's bad enough on a neonatal mouse, but on something only a few millimetres long!

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Could thymus lymphocytes become immuno-competent?

In 1965 Gus Nossal succeeded Sir Macfarlane Burnet as Director of the Walter and Eliza Hall Institute, in Melbourne. And he remembered his schoolboy friend.

Yes. Being an immunologist he had been interested in my work, and he invited me to the institute. I decided to go back to Australia partly because my wife, being Australian and having a sister there, would like to go back, and also because I realised that Gus would give me tremendous encouragement and a lot of facilities, many inbred strains to work on and so forth.

You haven't regretted that decision?

No, not at all. The Walter and Eliza Hall Institute is a magnificent institute and a great place to be. The infrastructure is stupendous and it is just wonderful that you can talk to any scientist, whether or not they are in your field. Gus is to be congratulated very much for having built an institute in such a way.

Even when I was there, in Burnet's day, it was a wonderful place to work in and very bright visitors would come out from Europe and America to work with him for a year or so, but it was really tiny in numbers of people – about the size of a small department of a modern institute or university. Gus's first idea was to build it up, and he did that very effectively.

It was still quite small when I first joined, but they built two floors and then one floor on top, and then finally you had a completely new building.

When you joined the Hall Institute in 1966, what did you embark on?

I was pleased to find that Gus had already chosen Graham Mitchell, who had just graduated in veterinary science from the University of Sydney with first-class honours, to be my first PhD student. We made it our first priority to see what sort of cells in the lymphoid and haematopoeitic system could restore immune responsiveness to thymectomised mice, either neonatally thymectomised or adult-thymectomised, irradiated and marrow-protected.

To explain: the effects of neonatal thymectomy can be reproduced in the adult, provided after adult thymectomy you get rid of all the lymphocytes that are present in the body, by irradiation, for example. If you give very high doses of irradiation you have to inject bone marrow, of course. But in the absence of the thymus, the stem cells coming from the marrow can no longer generate lymphocytes, at least thymus-dependent lymphocytes. So you could reproduce the effects of neonatal thymectomy in the adult, provided you followed that up with irradiation.

We decided to have a very, very good look at the types of lymphocytes which would restore immune responsiveness to immuno-incompetent mice. We used thoracic duct lymphocytes, removed from the lymph – and the spleen, of course, and bone marrow, thymus and lymph nodes – and studied their effects.

It was good, because thoracic duct cannulation was a very powerful technique which taught us a lot of things, for example that neonatal thymectomy – or adult thymectomy, irradiation and marrow protection – was associated with a tremendous diminution of the number of cells that could recirculate. Those recirculating cells were the cells which Gowans had shown to be immuno-competent, so again this showed that the thymus had something to do with the buildup of immuno-competence. But was it an effect of putting lymphocytes out to the circulation, or was it a humoral effect? It could have been an endocrine effect, as Medawar believed – that the thymus did not produce lymphocytes that went out in the circulation as competent cells but produced some kind of hormone which activated lymphocytes.

I remember talk of the 'thymic hormone', yes.

We really had to show whether thymus lymphocytes became immuno-competent. One of the experiments we did was to inject thymus lymphocytes which had been passaged twice in irradiated mice, together with antigen, to try and boost them up. But even those thymus lymphocytes which had been passaged with antigen into irradiated mice were unable to induce responsiveness. They were certainly not as good as lymph-node lymphocytes. But when these lymphocytes were given to adult-thymectomised, irradiated and marrow-protected mice, then the response was fantastic. In the presence of bone marrow, thymus lymphocytes were able to restore responsiveness, which in those days we measured by antibody production, on a plaque technique.

Yes, one thought immunity and antibody went together.

Yes, except for my skin-grafting, which depended on cellular immunity, not antibody. It had been shown that only lymphocytes (not antibody) could reject skin grafts. Thymus lymphocytes had never been shown to produce antibody, but when we showed that in mice that had been thymectomised and protected with bone marrow were able to restore antibody responses, we thought, 'That's fantastic.'

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Differentiating lymphocyte subsets

I said, 'Now we're going to show that thymus lymphocytes are the precursors of antibody-forming cells. We have to have a genetic marker.'

In those days they didn't have any known markers like theta or Thy-1, or CD4 or CD8 – they had none of the markers we have now. We had only the markers which were determined by histocompatibility genes, so we simply used an F1 hybrid thymus into a parental genotype recipient. In other words, we had a cross between two strains of mice, A and B, which can be distinguished by histocompatibility genes. The thymus came from an F1 and the recipient mouse was a thymectomised, irradiated, parental genotype strain mouse which had been protected again with parental genotype marrow. Into these mice we injected the F1 thymus, and we got a tremendous number of plaque-forming cells. And now, because we had antisera against both parents, we could determine whether the origin of the plaque-forming cells was thymus or bone marrow. The results were absolutely spectacular, no question about it. One plate was complete with plaques and the other plate had nothing.

We knew that one of these would tell us whether it was thymus lymphocytes that became antibody-forming, and I bet it would be thymus. It wasn't. It was bone marrow. But the thymus-derived cells were absolutely essential in order to allow the bone marrow-derived cells to make antibody, so there must have been some kind of interaction between thymus-derived cells and bone marrow-derived cells. We were able to publish those spectacular findings during 1967–68 in a very short statement in Nature; a paper which Burnet sent to the National Academy of Science, in the United States; and four papers in the Journal of Experimental Medicine.

That was the very first unequivocal demonstration, I think, that in mammalian systems you have two major subsets of lymphocytes, one of which is not derived from the thymus but becomes the precursors of antibody-forming cells. The other one is derived from the thymus but is needed to help in some way the antibody-forming cell precursors to become antibody-formers. That is somewhat similar in birds, which have both a bursa and a thymus.

I remember the bursa of Fabricius in the bird. Is the system the same as in mammals?

It is exactly the same, but the bursa doesn't exist in mammalian species. Its function, we know now, is taken over by bone marrow.

In hindsight it turns out very useful that 'bone marrow' begins with a 'b', so that you could talk of B cells in both birds and mammals.

Yes. We called them 'bone marrow-derived cells' and 'thymus-derived cells', which was quite a mouthful. For a long time I tried to work out short terms for them but it was Roitt, in London, that coined the term 'T and B cells'. I wish I had done that.

Roitt is a very good populariser of science who writes general textbooks for non-immunologists to understand, as well as immunologists.

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Eventual acceptance, but new complexities

To make two such major discoveries – the interaction of B and T lymphocytes in antibody production, and the role of the thymus itself – is quite an achievement.

Thank you. Actually, the concept of two cells interacting in the immune response was not accepted by the immunological community for a long time, for several years. First of all, Gowans had shown that the small recirculating lymphocyte can either produce antibody or reject skin grafts. To him the small lymphocyte was a type of cell and there need not have been two subsets with any difference between small lymphocytes that caused cellular immunity, or skin graft rejection, and those that became antibody-formers. It was well ingrained in the literature that one type of lymphocyte can do both.

Secondly, both T and B cells had to be specific for a given antigen. In fact, we proved that by using the antigen-suicide technique of Gordon Ada. We showed that we could suicide thymus cells with a specific antigen that was heavily radio-iodinated. So they had specificity of their own. And of course the bone marrow-derived cells had specificity because the antibody they formed was specific. So how could two very rare, clonally individuated cells ever meet each other in order to collaborate? People said, 'It is just not possible.'

How did you get over that? What was the next step in the investigation?

People repeated our work and eventually it came out to be absolutely true: there are two types of cells. Then came the antigen Thy-1, which someone had discovered in America – I wish I had used it before – and which was shown to be on some lymphocytes but not on others. In London, Mitcheson and Martin Raff used an anti-Thy-1 antibody to show that you could eliminate the type of lymphocyte that was thymus-derived and responsible for skin-graft rejection. It did not eliminate the other lymphocytes.

The other thing which came out was that one type of lymphocyte had immunoglobulin on its surface but not the other, and the one that had immunoglobulin on its surface was the bone marrow-derived one. As all this came together, people started believing in two subsets.

Among the interactions between T and B lymphocytes, essentially, a subset of T lymphocytes is important in its effect on the precursors of B lymphocytes.

Yes, that is correct. After this work, it was shown in America – not by us – that even T cells can be divided into two major subsets, CD4 and CD8. The first helps B cells form antibody and the other becomes the cytotoxic T cell, which kills off virus-infected cells and may be involved in graft rejection and so on. So even that is more complicated than people thought in the early days.

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Studying delayed-type hypersensitivity: could MHC restriction be shown in vivo?

Perhaps harking back to your early family history of tuberculosis, you started looking at the delayed-type hypersensitivity reaction.

Yes. That is a type of cellular immune response which we wanted to study in vivo. Many people at that time were studying cytotoxic T lymphocytes in vitro, in tissue culture. I was keen to study the interaction which takes place in vivo, so we decided to study delayed-type hypersensitivity (DTH, as I call it). With Matthew Vadas, another PhD student at the time, we devised a new technique for measuring DTH which relied on the radioactivity that you can measure in a particular site, in fact in the ear of a mouse. I didn't like to measure the thickness of the ear. It seemed to me that it is not so accurate, because it depends on the person who measures it. We devised a method of injecting a radioisotope in the ear so that the ear would have a certain amount of radioactivity and we could measure that effectively in a counter. It would not be dependent on the investigator.

Zinkernagel and Doherty had discovered the phenomenon of restriction by the major histocompatibility complex (MHC). This phenomenon had been demonstrated in vitro, and using this particular technique we were able to show that it also occurred in vivo in the transfer of delayed-type hypersensitivity. When you transfer DTH in vivo, you cannot transfer across an H2 barrier. You have to have H2 identity, which is similar to MHC restriction in vitro. I remember ringing Zinkernagel and speaking to Doherty about it in those days. They were very pleased that another system showed that they were correct.

If you had showed they were wrong, they wouldn't have been so happy! But if 'discoveries' are made in vitro, in cultured cells, it is very important to check them in an intact animal. Otherwise you may be led up the garden path.

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Immune-responsiveness genes: a masquerade

How did you go on from there?

At the time, 'immune-responsiveness genes' were in fashion. These were supposed to allow certain strains of mice to respond to certain types of antigen. For example, in the laboratory you could make synthetic antigens like a polyamino acid of glutamic acid, alanine, tyrosine, and you could show that certain strains would respond perfectly to these synthetic antigens but other strains would not respond at all.

When you say 'respond', do you mean produce antibody?

By delayed-type hypersensitivity or by antibody responses, by any criteria. The supposed immune-responsiveness genes were thought at one stage to be the receptor on T lymphocytes – the T-cell receptor for antigen – because they generally acted in the case of cell-mediated immunity or helping antibody formation. But our experiment with the transfer of delayed-type hypersensitivity showed that the immune-responsiveness genes were just the MHC-restricting element of Zinkernagel and Doherty – they were identical. And this turned out to be true.

So the term 'IR genes' has disappeared from the literature as a false conception. Sometimes things get simpler rather than more complicated! Since cellular immunity can be effective against intracellular parasites, did you think of investigating the idea that it might be effective against cells that had changed by becoming malignant?

Yes. Also, in my early work I had shown that neonatal thymectomy allows mice to develop cancers much more frequently than non-thymectomised mice. It was evident that this must be true to some extent but we did not investigate it in great detail. A lot of people in George Klein's laboratory, among others, were investigating tumour immunity, and it has been shown that there is such a thing as immuno-surveillance. But now not just T cells but NK cells, natural killer cells, are involved as part of the immune system.

That again shows the complexity of things we call lymphocytes, when NK cells are also morphologically just like the others. The cells that circulate in the lymph are really a pretty complicated lot.

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Where was that elusive T-cell receptor?

Where did you go after the work on DTH?

We did quite a lot of different things. It would be hard to single one out. I will tell you one of my great failures. We decided to look for the antigen receptor on T lymphocytes. It was clearly known that it was immunoglobulin on B cells which allowed B cells to make antibody. Several groups in the world thought, without actually knowing, that there was some kind of immunoglobulin molecule on T cells. In fact, in the Hall Institute, Jack Marchalonis coined the term 'IgT' for immunoglobulin T, being on T cells. I had never thought that was correct.

We really tried to find immunoglobulin on T cells, but I just wasn't able to show any evidence of it. With the help of a molecular biologist, David Kemp, we showed at the RNA level that there could not be any RNA expressed to make immunoglobulin on T cells. That was a negative result, which was written up in the Journal of Experimental Medicine. Trying to look for the T-cell receptor without much molecular biology knowledge, I failed completely to find it. The Americans found it!

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Why are we tolerant to our own tissues? More complexities

You have certainly done a wide range of different things. You did so much work in vivo, in the best experimental animal there is, the mouse – which now has been rendered vastly more valuable than ever before, even with SPF pure-line mice, by the possibility of making transgenic mice.

Yes. That, I think, rescued me from the T-cell receptor. When transgenic mice became available I immediately realised that this is the ideal system to study tolerance to 'self' – not tolerance, as Medawar and others had studied, to antigens introduced into the animal, but tolerance to one's own-self tissues. Why we are tolerant to self is still unresolved. When transgenic mice became available we were able to inject into the fertilised egg a gene which would make a gene product in a particular tissue. We chose a gene product against which we had an antibody, so that we could find its product. It is like having a marker for self, a self marker. All our work since 1986 or '87 has been done on transgenic mice and on self-tolerance. That has been very productive.

We have shown that there are several mechanisms of self-tolerance. In transgenic mice, both we and Zinkernagel have shown that some of it is due to ignorance. We have also shown that some tolerance is due not just to the self-reactive cells being eliminated in the thymus. If they escaped from the thymus they could be eliminated in the circulation as well, as a result of antigen driving the cells to elimination – now shown to be due to a death gene called Fas, or CD95. These studies have been productive but have led to more complications, involving signalling in the cells by various types of molecules. Other people studied CD95 as one which signals the death of the cells, but we have looked at signalling molecules with names like CD30, CD40 and so on, which are very important in the immune response – or in shutting it off. That has great potential for the study of autoimmunity – autoimmune diseases, for example – and perhaps also neoplasms. If, instead of shutting the cells off, we could activate them to kill cancer cells, that would be very useful.

How is the shutting off done in the thymus?

Probably the cells which encounter self antigens in the thymus are killed through some mechanism dependent on the avidity of the T-cell receptor for its target. If the T-cell receptor has a very high avidity, then the T cell is killed. If it has a low avidity, the cell can escape or it can respond to other antigens. We have worked on that, but it was not discovered by us.

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Contributing to the scientific community

Would you tell us about some of your activities in national and international bodies?

I served on the Academy's Council and Sectional Committees, and a few other things like that. Also, I was asked to serve on several international agencies.

For 3 or 4 years in the 1970s I was on the International Agency for Research on Cancer, whose headquarters are in Lyon, and I eventually became the President of the Scientific Council. It was interesting to see what sort of research is done in that field, and I benefited from that.

I served on WHO for quite a number of years, on eradicable diseases. WHO wanted to eradicate five parasitic diseases by the end of the millennium, including malaria, schistosomiasis and leprosy. It was very ambitious: I don't think malaria will be eradicated by the end of the millennium. I was one of several immunologists on that committee.

So it was strongly oriented towards vaccine production?

Definitely towards vaccination, yes. We were also trying to boost research by checking grant applications and deciding on various methods of attacking those diseases. I also served on WHO every now and then to summarise certain aspects of medical research – immunological research, in my case – writing books that the WHO put out on certain topics.

You did a term on the International Union of Immunological Societies, and you have regularly attended conferences on immunology, from way back in 1961 up to the present.

For many years, with three or four international trips each year. I have been quite active in that respect.

You have given us a very good picture of your career, Jacques. Thank you very much.

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Sir Otto Frankel was a geneticist by training, plant breeder by occupation, cytologist by inclination and genetic conservationist by acclaim. Apart from his personal research, Otto was a highly effective builder and leader of research groups, Socratic gadfly to the scientific establishment, and high prophet of the genetic resources conservation movement. His career in science was unusual in that his most widely acclaimed work was done after his official retirement.

Interviewed by Dr Max Blythe in 1993.

Contents

• Cursedly indpendent from a very early age
• The simple matter of becoming a geneticist
• An Austrian Jew's entry into the British Empire
• A Cambridge introduction to cytogenetics
• The New Zealand years: chance encounters, enduring benefits
• Promoting good science in the CSIRO
• Linking developmental genetics with plant physiology
• 'Let's call it Genetic Resources!'
• Genetic diversity in nature reserves
• Honest science for self-sustaining ecosystems

Cursedly independent from a very early age

Sir Otto Frankel, it's good to be able to talk today to someone who has made such a massive commitment and contribution to gene conservation. First, though, can I take you back to beginnings? You were born in Vienna in 1900, I believe.

I really don't remember much of my childhood, but I don't think either of my parents was an especially powerful influence on me. I went to school in Vienna, but after that I was interested to move away – not because there was any unkindness in my parental home but just because from my very early days I was rather an individualist and cursedly independent. I was also a naive young man, stupidly idealistic. I was anxious to go to Russia, with no idea that I wouldn't have lived very long, had I gone there. I became a Communist and didn't shed that until years later.

Similarly, my own stupidity made me take up agriculture. I had no idea of how to do any farming or produce anything, but after the First War the world was very hungry, especially central Europe, where I was studying chemistry in Munich. Suddenly it occurred to me, 'The world is hungry. Why am I wasting my time on chemistry? I should be doing agriculture.' And so I started studying agriculture in another university, in the western Germany town of Giessen.

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The simple matter of becoming a geneticist

Then I got so tired of studying that I gave it up altogether, until an old aunt of mine (a farmer on whose farm I was working) said to me, 'Otto, you're not an idiot. You're quite an intelligent young man. Don't waste your life like this. Go and study properly.' With her financial help I went to the Agricultural University of Berlin, which had a very high scientific reputation.

In my second year there I heard Erwin Baur, a very distinguished German geneticist, lecturing about genetics – something which was completely new to me; I'd never even heard of it before. I was fascinated. I thought, 'Well, this is it!' After a few lectures I went to Baur and said, 'Professor, can I do a thesis with you?' He said yes, when could I start? 'Next week.' He fixed a time for me to go and see him about it; I started my thesis; and I became a geneticist. It was as simple as that.

I worked on the snapdragon, antirrhinum, which Baur had made quite a famous object. It is now very much in the forefront in molecular work and people have done wonderful work on it – but still based on the background information provided by the work of my generation. Without that part of my life I don't know what would have become of me. My doctoral thesis included the first review on linkage in plants, where there was already quite a literature, which was then published in the German genetical journal as a major contribution.

What happened after that?

Well, although I was Dr Frankel by then, I had nowhere to go, nothing to do in life. It was very hard. One of my father's clients, an Austrian baron, had a large sugar factory and farms producing sugar beet in Slovakia, very close to the Austrian border, and my father arranged that I would be able to work there without pay. So that is where I went.

I was still far too young to apply for academic posts, and anyway I hadn't contemplated that. At the back of my mind I wanted to produce food. And Russia was still at the back of my mind, but in my naivety I had no idea how to do anything about it. I just sat there, not learning anything. But I did gain a little bit of experience; there was no-one who knew more than I did.

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An Austrian Jew's entry into the British Empire

Were you hindered at that time by your Jewish background?

I'm sure I was. I really didn't have any openings. Working in Germany seemed unthinkable, partly because of the Jewish background.

But that background became very useful. (I hope you won't mind if I tell you another personal story.) One day I got a letter inviting me to come to London in connection with a post in Palestine as research officer in a team in which the Empire Marketing Board was involved. This Board was an imperialist move after the First World War to establish connections with the Empire – commercial connections on behalf of the motherland, rather than on behalf of the colonies. But the Board also had a committee to establish research schemes in the dominions and colonies. The emphasis was on mineral deficiencies, especially minor element deficiencies, which were quite fashionable at the time, being very important in animal production. A number of minor elements had been discovered, but the question was where else they existed.

The chairman of the research committee was Major Elliott, DSc, a member of parliament who had a close affiliation with the Rowett Research Institute at Aberdeen. And the Institute's director, Dr John Boyd Orr – later Sir John and then Lord Boyd Orr, the first director-general of FAO – interviewed me, very informally, in London.

In fact, my first dinner in London was in the House of Commons, where Walter Elliott had invited my cousin, Lewis Namier – who was already a distinguished modern historian with the chair at Manchester, and later became Sir Lewis. At that time he was the political secretary to Weizmann, the founder of Zionism. A group of Conservative politicians affiliated to Lord Balfour (of the Balfour Declaration for the Jewish national home) was attempting to assist the Zionist development, and Elliott and Balfour's niece, Mrs Blanche Dugdale, were the centre of this group. This research scheme on which I was engaged, then, was a cooperative venture between the Empire Marketing Board and the Zionist organisation, but actually administered and run by the Colonial Office.

So I was appointed by the Colonial Office and went to Palestine with a young Aberdeen man from the Rowett Institute, John Crichton, with whom I became firm friends and who taught me a lot of very colloquial English, with a great many swear-words! And that was my entry into the British Empire.

An amazing experience. How long were you in Palestine?

I stayed there for 9 or 10 months. But I was a plant geneticist and breeder, and animal nutrition really wasn't my field. I had been promised that if I didn't like it in Palestine they would bring me back to England and help me find a new job in my field, and so I came back – first to London and later to Cambridge.

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A Cambridge introduction to cytogenetics

You were in Cambridge for nearly a year then, in the '20s. What were you engaged upon? It certainly impressed people.

I was working in the Plant Breeding Institute (which was part of the university) with A E Watkins, a very good wheat cytogeneticist and evolutionist. I had worked on wheat in Czechoslovakia, but it was Watkins who brought me into the cytogenetics of it. I learned a lot from him. He was a nice chap, who among other thing introduced me to English literature – he made me read Jane Austen and a lot more which was quite unknown to me, and seminal.

Watkins was very interested in establishing a collection of land races. He was obtaining material, in Cambridge, through the British Board of Trade. Its officials in different parts of the world, including Mediterranean and Near Eastern countries, went into markets and collected samples of seed lots that were for sale. Watkins generously shared his samples with me, and later when I came to New Zealand I had about 3000 in my collection.

Really, the science of it was based on the work of the remarkable Nikolay Vavilov, a Russian explorer and plant geneticist, plant geographer and collector. And 10 years later, in 1935, I spent a week with him in Leningrad. That link with Watkins and Vavilov in my Cambridge days influenced me greatly throughout my life. The cytogenetics of wheat became my field in New Zealand for many years.

I think your move to New Zealand had something to do with Biffen, who had been your first contact in Cambridge. Is that right?

Biffen was the Professor, but I saw him only once when I arrived and, of course, paid my respects. I never saw him again until he brought me the telegram which started me on the way to New Zealand.

That telegram is another story of the utmost chance. It came from John Orr, who had crossed Canada by train (as one did in those days) with Ernest Marsden – an English atomic physicist who had worked under Rutherford and had come to New Zealand as an administrator of education, I think. When New Zealand set up a Department of Scientific and Industrial Research (DSIR), as everyone did at about that time, Marsden became its head. So Marsden told Orr that having established a dairy research institute and appointed a cereal chemist, he was now establishing a wheat research institute and looking for a wheat breeder. It seems to me that Orr pricked up his ears at this and said, 'I have got him for you.' The result was his telegram to Biffen, who asked me if I was interested. Of course I was. And that greatly important New Zealand chapter of my life lasted for 23 years or so.

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The New Zealand years: chance encounters, enduring benefits

Tell me about those New Zealand years, Otto. You went to a government-type job?

Very much, because I was appointed to the staff of the Wheat Research Institute under the DSIR. It was interesting enough, and I could do other things as well. I became very good friends with my director, F W Hildendorf, who was a leading man in agricultural science development in New Zealand and also Professor of Botany at Lincoln College, in the University of New Zealand. When he died, I took over from him.

During those DSIR years I did a lot of work, not only on wheat but also in the cytology of quite a different plant. In fact, that was the work for which I was made a Fellow of the Royal Society, in 1953.

The main outcome of your New Zealand work would have been the wheat strains you created, I suppose.

Oh well, I did some very difficult cytogenetic work there which was published in a series of scientific papers. But in my mind, as I look back at my life, none of this was really outstanding or as important as the work on genetic resources, most of which came after I formally retired. I look on the work we did on genetic resources and the genetics of conservation as my main contribution, the thing my name is reasonably well known for.

Wasn't it in New Zealand that you married your present wife?

My second wife, yes. I always think, you know, that through that chance interview on the railway train between Orr and Marsden she got a wonderful husband!

But in 1951 you moved to Australia. Did that also happen by chance?

In effect, yes. I had never seriously thought of looking for a job until quite late in my stay in New Zealand, when I had a row with the chairman of the Public Service Board (a personal friend). I wanted to appoint to a job a young Australian who wanted to continue his Australian salary level until the equivalent salary under our scale caught up to it. But this would have meant circumventing our very fixed salary system by a few hundred pounds – a trivial amount – and the Commissioner refused to let me do it. I thought his reason was plainly stupid, and as I walked out I said to a man from our head office, 'The next job that's going! I mean it.'

Then, one day, a colleague asked whether I had looked at the Australian advertisement that was on the noticeboard. When I admitted that I never looked there, he said, 'You ought to. You are the only one here who could take an interest in that one.' Well, I took it off the board, wrote a letter of interest and named referees, and was invited to come to Melbourne. Had this man not spoken to me about the job, oddly enough, I would have missed it.

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Promoting good science in the CSIRO

You came to the CSIRO, where you remained.

Yes, and straight away I was enormously impressed. It was a different, larger world.

The post was as head of the largest division in CSIRO, the Division of Plant Industry, and the selection committee consisted of a number of university professors (several of whom became my good friends later) and some heads in CSIRO. I wasn't terribly keen, actually, when I was taken to Canberra and saw so many difficulties there. The main difficulty was, as I told the chairman of CSIRO, Sir Ian Clunies Ross, the next day, that I felt terribly ignorant. I had previous experience of hardly any field in the Division, and I told him I just felt so inadequate to fill this post that I really couldn't accept it if it were offered. But that impressed him enormously, as I realised later on, because I was the opposite of self-advertising.

As we talked, I kept on asking what they had done to find a more suitable person. I said, 'How about so-and-so?' and that impressed them even more. I couldn't understand why they didn't appoint the man who was acting chief, a distinguished ecologist and pasture expert, but Clunies Ross said simply, 'He would make it into a pasture research place, but we want very much more than that. We want good science.' So I was offered the job, and good science they got – in the end, for him perhaps a bit too much science and not enough agriculture. We went on to have a close, friendly relationship, however, and he played a big role in my life.

In the late '50s your administrative work grew, when you were on the Executive for four years. Did you enjoy that role?

No, I didn't, although really I was quite a good administrator. When it was offered to me, I accepted it because the large divisions were being disrupted and made into smaller ones but Plant Industry as it had evolved was such a good place, so productive, and had assembled such a very good staff that I didn't want it disrupted. I wanted to appoint my successor and see that he was safeguarded. With that done, the place got better and better rather than weaker and weaker as I had feared it might. It's a highly distinguished institution now, of considerable world reputation. I know I am given a lot of credit for this, but I did at least sow the seeds to make the beginning possible.

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Linking developmental genetics with plant physiology

Clearly that saved the Division. With such a big administrative role, though, did you have a chance to do bench work?

I always had a small group, usually of two people, working with me on the genetics work which I had started and which ran through my life until the '70s, when it turned to physiology rather than cytogenetics.

I just thought you looked at a lot of chromosomes all the time.

No, not in this work. This was developmental genetics, work on the genetics and development of the wheat flower, and it was very difficult. We had discovered two different systems of genes which are responsible for the development of the wheat flower and can be found in a group of mutants called speltoids. They are similar to spelt, which is a species of the Triticeae and is closely related to our bread wheats. It is distinguished by a large, complex mutation involving several genes.

If this mutation is present, it can be transferred from mutants to bread wheat. When it is there, a different system of flower formation is revealed once the gene responsible for non-speltoidy is removed, and it is this secondary system which I studied throughout the years, using physiological methods. In Plant Industry we had developed a phytotron – that was one of my major achievements in Australia – and so I could use it as a place to do the physiological work. The series of papers which resulted didn't end till long after I retired. (I stayed on in the Division of Plant Industry doing this work, in addition to the international work on genetic resources.)

Your work changed the production of wheat and flour. Did it have an economic outcome?

No, not really. But it was of considerable academic interest.

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'Let's call it Genetic Resources!'

How did you eventually become involved in working on genetic resources?

Well, there were two components of the curious way that came about. In the 1960s I was asked to represent Australia on the international scene. As it happened, Ledyard Stebbins – a very distinguished friend of mine and a well-known American evolutionist and taxonomist – took a leading part in the earliest stages of the International Biological Programme, and because he thought I should become interested in what became known as gene pools, he bullied me into taking over that field in the new IBP. So this is how it all started.

To make a long story short, it got me into the Food and Agriculture Organisation as a temporary consultant to advise on what they ought to be doing about genetic resources, as we later called it. I was one day sitting in my office in FAO when in came Hermann Kuckuck, a German agricultural scientist, younger than I, whom I had known a little. He told me a very interesting story, that extremely important genetic resources in Iran and in Abyssinia (Ethiopia) were disappearing. This was quite new to me and it made an enormous impact, because I had been interested in plant collecting and plant collections for many years, almost from my student days. As I then followed this up and got information about land races – the old peasant varieties which had been selected by farmers and also, very largely, by natural selection, and which had been the real resources of plant breeders ever since plant breeding started – so my interest in genetic conservation grew.

An international meeting of experts in Rome, which I organised, was the formal beginning of the whole campaign to assemble and to salvage the genetic wealth that had been accumulated in different parts of the world where agriculture had evolved. That was really the beginning of it for me, and the beginning of the movement to save genetic resources. At that 1967 meeting, I was talking to my colleague Erna Bennett, who had come to FAO to help with the final phases of organising this conference, about what we were going to call this. And together we said, 'Genetic Resources!' That's where the name comes from, and that is how we started the campaign against the loss of plant and animal species. And I'm still very much involved.

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Genetic diversity in nature reserves

What was the other component of your involvement in genetic resources work?

That was an invitation to give a Macleay Memorial Lecture, in a series which had been established by the Linnean Society of New South Wales. I wasn't actually very intrigued, but such distinguished people had given that lecture before that I thought it would be pretty immodest if I were to refuse.

I was going to talk about what I had done before, but whilst I was thinking about genetic conservation it suddenly occurred to me, 'We plant breeders have all these land races and wild relatives of crops as our resources, and we bring them together, we cross them and we select. But what happens in nature reserves?' God would have to do it, and I couldn't see God coming down and making all these crosses! How would Nature have the diversity to select from for natural selection to occur when the environment changed – say if the climate hotted up or got dry or wet? As that sank in to this dumbskull head, I thought, 'This is quite something. There must be genetic diversity in nature reserves.' And this is why the Macleay Lecture is, to a degree, quite seminal. People hadn't talked about this. Perhaps everyone took it for granted, but no-one had emphasised it. And now it's become a bandwagon and it's being overemphasised.

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Honest science for self-sustaining ecosystems

Tell me about the book you are working on at present, Otto.

It is like a sequel to the one that came out of the contributions to the 1967 conference, which became really the first bible of genetic conservation. In writing The Conservation of Plant Biodiversity, Dr Tony Brown (A H D Brown), Dr Jeremy Burdon and I are trying to establish a scientific balance. Biodiversity has become a catchword of the first order, but in this book one of the subjects we are dealing with is justification for species conservation.

You see, conservation takes two major forms. One is the conservation of communities, which we strongly advocate, and the other is conservation of individual species, whether of particular significance or not. Many people, including very distinguished American biologists, now say that everything must be preserved – simply, as one of them said to me the other day, 'Because it's there.' To me that makes little sense, so in this book we discuss various aspects of species conservation, among many other things, in a scientific way.

The three of us are firm that the important issue is the conservation of natural areas, although we have some differences about letting things die out. I am quite cheerful about that; I see no reason for attempting to preserve everything just because it's there. There must be a degree of selectivity. But I am all for preserving as many ecosystems as we can contrive to. I don't think we are winning in that battle, however, because of the increase in human populations everywhere. We can't win until we ourselves stop growing. This is not just about gene pools but about communities – complex communities which can sustain themselves and develop through natural selection.

Will your book be published soon?

We have a contract with Cambridge University Press for termination at the end of this year. Each of us has three chapters to write. I have written mine, and revised and re-revised them, but my colleagues are not as far advanced because they have a lot of other things to do. Besides their own research, they have to earn their keep, and their jobs have become very arduous: CSIRO has become burdened with an enormous amount of administration, and they have got to write reports and help find money for their work. They have become subject to a change which I personally cannot approve of, but which is a reality. They are very busy people.

In addition, we need an introductory chapter which sets the scene. I have written much of this, but when they have done a bit more work I will have to write some more introduction to refer to their chapters. And then there will be a summing up, in which we have to stress what we believe is important in the conservation of plant biodiversity. There we have to come out as honest scientists.

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Professor Mervyn Paterson is a geophysicist who has led Australian research into rock mechanics and pioneered instrument development over the last fifty years. He was born in South Australia in 1925 into a family of wheat farmers. He attended Adelaide Technical High School, then The University of Adelaide from 1941 to 1943.

He began his career at the CSIR Division of Aeronautics working on the physics of metal fatigue, a foundation which shaped his entire career. He received a PhD from The University of Cambridge in the UK on x-ray diffraction effects of deformation metals, and pursued postdoctoral studies in Chicago in the USA. He returned to work at the newly-named CSIRO, but soon moved to the Australian National University, where he stayed for 31 years in the Research School of Earth Sciences. During this time he developed instruments to test rock deformation, which subsequently led to a 'second career' as owner and manager of Paterson Instruments P/L, a company specialising in building scientific instruments.

Interviewed by Professor Kurt Lambeck in 2006.

Contents

• Studying rock deformation
• The importance of crustal rocks
• Influences in rock deformation studies
• Early life and family background
• Widening horizons at high school
• From university metallurgy to CSIR research
• A Cambridge PhD in material science
• Chicago: the postdoctoral student meets his wife to be
• Transition from CSIRO to the ANU
• Moving firmly into experimental rock deformation
• A new career in equipment building
• Whither rock deformation studies?

Studying rock deformation

I think it is fair to say that Mervyn Silas Paterson is Australia's leader in research into rock mechanics under a range of laboratory and geological conditions. And over some 50 years he has pioneered apparatus developments that have been adopted world wide and are still directing the research of this field.

He was first appointed to the Australian National University in 1953, becoming a professor in 1987 – when professors were still few and far between and the title actually meant something. He was elected to the Australian Academy of Science in 1972, and during his career he has received a number of international and national awards in recognition of his work, perhaps most notably the Walter Bucher Award of the American Geophysical Union.

Mervyn, it is appropriate that we start with your science. I recognise that it is futile to try to summarise your career in a few minutes, and the details would probably be better set out elsewhere. But perhaps you can give us an overview of what you have been doing all these years, and an idea of how it relates to geological research in general.

I really started as a metallurgist and later got into material science, as it is called nowadays. Coming to the ANU in 1953 represented a big change in direction for me, because that was when I got into experimental rock deformation studies.

We were trying to understand how rocks deform – to understand geological processes, because during mountain building rocks get really screwed up and twisted and bent. The object in the lab is to try to understand what the processes are, and what sort of strengths rocks have. How strong is a rock under geological conditions? What sort of forces do you have to apply to deform it?

So that is the sort of thing that has kept me busy over the years.

If you go to a road cutting and look at how the rocks there have been folded and faulted, how do you relate this back to your laboratory experiments?

Well, you have to imagine what the conditions were at the time when those rocks got deformed in that way, when they were deeply buried in the Earth, perhaps even at the bottom of the Himalayas. A rock normally is rather brittle – if you try to bend it at atmospheric conditions, it just breaks. So to do our experiments on plastic deformation of rocks we have got to find high-temperature, high-pressure conditions, say thousands of atmospheres of pressure and 1000 degrees in temperature. You can bend a rock just like a piece of copper if you have the right conditions.

And where does that place you in the Earth?

Perhaps halfway down in the crust, 15 to 20 kilometres. But once you are under those conditions you are studying processes that are no longer so pressure dependent, and so you can apply the results in thinking about deformations deeper in the Earth.

How do you extrapolate from your laboratory work to the geological environment?

That's a very tricky one, because we don't have the geological time in which to do experiments. That is where the detailed study of the structure of the rocks and the specimens and the crystals in them becomes very important, because we want to understand the mechanism of the deformation. There are various ways in which a rock could deform at the atomic or crystal structure level, and we need to look in the microscope for evidence of what processes occured in the crystals of the rock. That tells us something about what particular deformation mechanisms were effective.

Then we can go to rocks in the field, do similar studies there, and try to come to conclusions as to what the mechanisms of deformation were. If we can persuade ourselves that the same mechanisms were operative in geological deformation as were operative in our experiments, we are entitled to extrapolate our very short-term experiments – hours or days – through to geological time of millions of years.

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The importance of crustal rocks

During your career there has been a major revolution in the earth sciences as the plate tectonics hypothesis has developed. How has your own science been influenced? Has it made the outcomes of what you are doing more relevant?

I had got into this field about a decade before the plate tectonics revolution so it didn't greatly influence what we were doing. It has been extremely important for the context of geological studies, which I suppose has impacts on the laboratory work, but plate tectonics is really very large scale and we work at the intermediate and small scales.

I guess one consequence has been to move the emphasis in the earth sciences away from the crust into the mantle – to look at convection, for example, the dynamics of the mantle. Do you ever think of trying to do deformation studies at the greater depths and temperatures that are more representative of the mantle?

Well, there is no problem about the temperatures, because those we work at are at least the same as in the upper mantle. And the pressures are not a great factor in extrapolating down to greater depths, because the plastic deformation processes that we are looking at are not so pressure sensitive. It is exceedingly difficult to do experiments at the higher pressures. People are doing that now: a group in America and another one in Bayreuth, Germany, are doing experiments at much higher pressures. But they are much cruder experiments. I prefer to stay in a regime in which I can do more precise measurements.

I guess there is a question of choosing materials. If you are working at crustal depths you are looking at crustal materials; to work at mantle depths you have to look at mantle materials. Initially you have been working mainly on calcite, and later on quartz. How pertinent are these to understanding the Earth?

They are very pertinent in the crust. And I must take issue with the notion that all the important questions are down in the mantle. Most of the geology that we see is in the crust so I am a firm believer in the importance of working on crustal rocks. But we have also done a lot of work on olivine rich rocks, and olivine is the principal constituent in the upper mantle so we haven't neglected that. David Kohlstedt, from Minneapolis, spent a sabbatical here at a time when we had already got into working on olivine rich rocks, through work by Pram Chopra. Since then Dave Kohlstedt – using one of our machines – has taken up this sort of work in a big way in Minneapolis, which is now the centre of work on olivine rich rocks.

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Influences in rock deformation studies

Over the years you have had a tremendous impact on students, staff and visitors, many of whom have gone on to positions of distinction and influence in Australia and overseas and have shaped the discipline of rock deformation studies. To what do you attribute this influence, and what lessons can we draw?

That's a bit hard to answer [chuckle] (assuming that I have had much impact on people). Comments from one or two of my colleagues suggest that they have learned something about how to be meticulous in thinking and in examining and analysing problems. I think it is a matter of attitude and of a disciplined analytical approach.

And of your ability to marry the instrumental development with the science?

That's all part of the same approach. When you want to attack a problem you have to develop the procedures for attacking it, and if you are an experimenter that means developing experimental equipment.

I think it is fair to say that your research work is characterised by asking critical questions and then designing experiments in search of the answers. Can you give us an example or two of that?

Well, take for example the crystallographic preferred orientation in marble. Marble is made up of crystals of calcite and in many cases the crystallographic axes are lined up, they're not just random. It is thought that they are lined up during the deformation. In the lab we can put pieces of marble – calcite aggregates – into the apparatus and deform them, and then use X-rays and so forth to measure the preferred orientation. In that way you can actually relate the preferred orientations that you see in a marble specimen to the sort of deformation that it has had. That enables you to go into the field, pick up a piece of marble and say, 'That's probably been deformed in this way.'

What have been your most important contributions so far in this field?

I like to think my work on quartz has been important. Quartz is really one of the most difficult of materials to deform and I don't think we understand it altogether yet, but I think I have made a contribution there. Another one that comes to mind is some work we did very early on with Barry Raleigh, on deformation of serpentinite at high temperature. That work brought up ideas about how deep-seated earthquakes may occur and has had quite a bit of attention over the years.

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Early life and family background

Going back in time, Mervyn: you grew up in a 'one-horse town' in South Australia. In fact, you had to share a horse in order to get to school. Have any aspects of that early environment shaped your subsequent life?

I grew up in Booleroo Centre, a town about 300 kilometres north of Adelaide in a marginal wheat farming area – very marginal, I think. We lived right on Goyder's Line, which was supposed to be the limit to where you could grow wheat.

My forebears had gone up there in about 1875 when the area first opened up for wheat farming. So I came from a farming background and grew up on a wheat farm. The early 1930s were fairly tough years, however, when we had a lot of droughts and dust storms as well as economic depression, so things weren't very easy on the farm. But it was a great background in a way.

I went to local primary schools: small country schools with about 14 children in each of them, all classes in the one room with one teacher. The older kids got told off to teach the younger ones, and it was a cooperative venture. That's not a bad way to start off, actually. It teaches you a lot of self-reliance and so forth.

We had to travel quite a long way, though, and yes, I used to go to school on a horse. In fact, during the Depression our farming went back to the use of horses.

We escaped from going bankrupt in 1936 and moved down to Adelaide Hills, where I did my last year and a half of primary schooling at a very small, one-room school with a remarkable teacher, Max Wardrowski. He had a university degree, which was exceptional – all the teachers before that had been young women who were more or less just out of high school with a year's teacher college training. But because Max had a degree I became aware that universities existed and that a person could have an academic career.

Even earlier, I understand, your background includes the very foundation of Adelaide. Have I got that story correct?

Yes, one of my great-grandmothers came out in the first year of the colony. All my forebears were in South Australia by the 1850s. Half of them were from Scotland, a quarter from Wales and the rest from England, especially Somerset I think, with a great-great-grandmother from Ireland somewhere along the line. They were all farmers. My father used to have stories about the amount of cider that the 'Somerset' lot would drink during the summer.

I gather that your forebears were fairly staunch Protestants. Did that rub off on you and have any influence on your life?

[chuckle] I wonder. I think it's deep down. I think there were Cornish Methodist connections going right back to John Wesley himself. Certainly I grew up in a context of Protestantism. My father was a Methodist lay preacher, so we used to be regular churchgoers. I'm not particularly a churchgoer these days and I suppose I have become a bit agnostic, but I don't regret that as a background.

Mervyn, in reading obituaries of great scientists I am struck by how often their background has been very similar to yours – they have come from small towns, gone to small schools, had a not particularly academic environment. What is it about such a background that produces outstanding people?

I think one of the important things is self-reliance. When you come from such circumstances you do have to learn to make your own way in many respects.

I believe that even today you still walk to and from work. Do you think that mothers have to stop dropping their children at school and picking them up, and let them walk five miles?

[chuckle] Well, I don't think walking five miles does them any harm! It probably does them quite a bit of good.

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Widening horizons at high school

Did you go on to a local high school?

There was no high school in our area so I went away to the Adelaide Technical High School. It was a select entry school with emphasis either on engineering or on commercial activities. (Max Wardrowski had probably brought our attention to its existence.)

When I was about to start at high school there was a polio epidemic so school was a little late starting that year. We had correspondence lessons for a while.

The school was very good, with high standards, and I had very good teachers there. For example, my teacher in the first year was Max Bone, who was subsequently Director of Technical Education in the South Australian government. Another outstanding teacher was the chemistry teacher, Dougal Slee. I learnt a great deal from him.

I suppose I gradually developed an interest in pursuing further studies. The high school didn't teach any foreign languages, so when I got the idea that I might be able to go to university I had to go and do Intermediate French at night school while I was doing my Leaving. I somehow managed to pass it.

That must have stood you in very good stead later on, because your French connections are well known. At some stage during high school you had to make a choice, I understand, between going to university and joining your father as a farmer.

Yes. During the Intermediate stage he came to me one day and said he was interested in a new property down in the south-east of South Australia, a bigger property which would have been more than he could manage. Was I interested in going in with him in that? Well, I thought that I would like to do another year at high school yet [chuckle] and passed that up. I guess by that time I was getting doubtful about pursuing a farming lifestyle anyway. I had seen plenty of how tough a life it was, the financial stringency and so forth.

So that was a turning point. I guess I oriented myself to a non-farming life from then on.

I guess ignorance was bliss – you didn't realise how tough and difficult an academic life could be. You have no regrets about not becoming a farmer?

No regrets. I did think about going on to agricultural science, because I still was very interested in that sort of thing. My father had been very active in the local agricultural bureaux that used to keep farmers up to date with the latest ideas about farming. But one of my teachers, Stan Tiver, had a son who had just finished agricultural science and couldn't get a job. (This was just before World War II.) Well, considering all the unemployment of the '30s, the really important thing about a career was to be able to get a job! That ruled out agricultural science.

Dougal Slee had inspired me very much in chemistry and I thought that would be a good field to go into, but doing an academic chemistry degree didn't seem to be the avenue to a job either. So I did the next best thing and chose metallurgy, which involved plenty of chemistry and also had the possibility of a job at the end of it. And on the basis of that decision I entered the university.

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From university metallurgy to CSIR research

As I understand it, when you went to Adelaide University you were remarkably young by today's standards. What are your memories of university life?

[chuckle] I was just 16 when I started. I didn't take much part in university life in those days, except going to lectures and studying. It was very much a nose to the grindstone period. I used to do all right at it, though.

I started at the university in 1941. The end of that year was Pearl Harbor and the fall of Singapore, and the first vacation in 1942 we actually spent digging trenches in the grounds of Adelaide University because the Japs were coming down what were then the East Indian islands at an alarming rate. So the university engineering courses went into four terms a year, and the university shortened the course and gave an interim Bachelor of Science (Engineering) which I finished in '43.

Were you called up for military service?

I was called up, examined and pronounced A1, but as a metallurgy student I was sent back to the university because that was a reserved occupation. A metallurgist was supposed to be making cartridge shells and things, and wasn't allowed to join the Army. (I didn't protest about that.)

After university you joined the Aeronautical Laboratories at Fisherman's Bend, Melbourne, where you worked on the physics of metal fatigue. I understand that that was quite a remarkable place and going there shaped your subsequent career.

Yes, that was a real turning point. Our university course had been what one might call fire and smoke metallurgy – all smelting and ore dressing, extraction metallurgy, with practically nothing in the way of studies on the properties of metals. But in my last year at the university I happened to pick up in the bookshop a newly published book by C S Barrett called Structure of Metals, and that was an absolute revelation to me: I'd never heard about crystal structures in metals in my metallurgy course. It was an inspiration, and was partly the reason I moved over into the physical metallurgy field.

This move resulted also from having second thoughts about pursuing a primary metallurgy course. Professor Gartrell had lined me up a job at the Mt Lyell copper mine, doing studies on flotation of copper ores – which was right up my street, the thing that interested me most in metallurgy. But I got cold feet about spending the rest of my life in a mining town at that point, and Mt Lyell was about as remote a place as you can imagine, on the west coast of Tasmania.

Then I became aware, probably from an advertisement, of an assistant research officer post at Fisherman's Bend in the CSIR Division of Aeronautics, and I went there in the last year of the war. The lab had been going for about four years, I think. There was a remarkable galaxy of talent at that place, a real research environment, with people like George Batchelor and Alan Townsend.

They both went on to Cambridge and to great things.

Yes, very well known in the field of turbulence later on. There were a lot of people in mathematics; a lot of students of Tom Cherry, from Melbourne University, were there. Oh, it was a very stimulating place.

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A Cambridge PhD in material science

For your PhD you went to the Cavendish, in Cambridge. Did that mean you had to leave your position at CSIR?

Only temporarily. I went to Cambridge on an Angas Engineering Scholarship established by the Angas family of Angaston in South Australia – who had actually brought my great-grandfather out in the 1850s.

And that closed the circle.

Yes. The scholarship, however, still paid about the same amount of money as it did when it was established in the 1890s, and it was totally inadequate support. So I also got a CSIR studentship to make the Angas up to the value of their studentship for me. That was very good.

Can you tell us anything about your time in Cambridge? Did any particular events or individuals shape your future?

Going to Cambridge was a new step, and for me a step into the university life. While I was at Adelaide University I lived with my grandmother and life was mostly just studying. But in Cambridge one came in contact with all sorts of people and enjoyed the college life. For example, a couple of my very close friends there were Sahkar – subsequently the editor of the Times in India – and Abuticknama, who became Vice-Chancellor of the university in Colombo, Sri Lanka.

I went to the Cavendish to work with Orowan. I probably got to know about him from Walter Boas (a one-time Fellow of our Academy here) who had been a fellow student of Orowan's in the Technical High School in Challottenberg, Berlin, in the 1920s – a great period in German physics.

There was a very interesting group of people around Orowan in the Cavendish at that time. One of the people I used to share an office with was Rodney Hill, for example, who was just finishing his PhD in plasticity theory. Norman Petch was another interesting person. Robert Cahn is very well known in material science nowadays; he also shared that office. [chuckle] It was a very stimulating time, and I very much enjoyed living in Clare College and associating with the people there.

When you went to Cambridge was there again a change in research direction?

Going there was essentially a new orientation for me. It provided me with a new direction, working in X-ray diffraction effects of deformation in metals.

Actually, I didn't go there with a thesis topic in mind, and initially Orowan said I should work on the mosaic structure of crystals. Well, I spent about six months bashing my head on this and trying to understand the dynamical theory of X-ray diffraction and so forth, but I wasn't really getting anywhere. So he suggested one day that I look at the X-ray line bordering in copper deformed at liquid nitrogen temperature. I picked that up and I managed to extend it out to a PhD.

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Chicago: the postdoctoral student meets his wife to be

After Cambridge, then, you returned to Australia.

Yes, I came back to Fisherman's Bend for a year. While I was in Cambridge, CSIR got its O and became CSIRO, and when I came back it was to the Department of Supply, because CSIR hadn't been seen as secure enough for aeronautical research. It was all right during the war: we didn't even have a guard on the gate in those days!

Afterwards you went to Chicago for postdoctoral studies. Why did you pick Chicago? What was going on there?

That came about by chance. I think Orowan had received information about the postdocs going in Chicago and mentioned me, and I said yes, I might be interested in that. So he wrote off and they offered me a postdoc. But I thought I was committed to go back to Fisherman's Bend at that time and didn't take it up immediately. I was already in the boat on my way back when I got a cable from Fisherman's Bend saying they would okay me to go to Chicago. [chuckle] Well, I held them to that and a year later I went.

In Chicago I actually worked for a year with C S Barrett, the man whose book had inspired me earlier on.

The CSIRO were quite content to encourage you down this path?

Yes, they agreed to it. They had been cooperative all along.

Not only did the Chicago year change your science, but I believe it was in that big and foreign city that you met your wife to be.

Yes, that's another opening-out of one's life. I was a Protestant colonial [chuckle] from remote parts of the Earth, she was a Hungarian Catholic with all the cultural background of Europe, including food.

So this was the civilisation of Mervyn, was it?

Oh yes, I'm sure she'd like the idea that she had civilised me.

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Transition from CSIRO to the ANU

From Chicago where did you go?

I went back to Fisherman's Bend. In a sense I spent about eight years at Fisherman's Bend, but for half of that I managed to be overseas.

What did you work on this time?

I started work on the more fundamental aspects of fatigue in metals. I was looking at reverse deformation effects, studying single crystals and pulling them back and forth, looking at the changes – defects and so on – in them. Actually, I spent much of that time perfecting a machine which I later brought up to the ANU and continued to use here. (My first research student did his PhD on that machine.)

We come now to your time at the ANU. You have been at this university for longer, possibly, than anybody else.

Except Frank Fenner! [chuckle]

Would you care to make any comments about what you have seen over this period and perhaps how science was done in the early days compared with today?

There has certainly been a lot of change over the years. Part of it is associated with the increasing size of the place. In the early days one knew everybody – I even knew all the people in Pacific Studies and Social Sciences, whereas I don't know anybody over there now. One was linked in more broadly to the university community in those days.

Finance was never really a problem: the university was quite well funded, and funded in the institutional way so we didn't have to spend our time writing grant proposals and so forth. And we were starting off in new fields, largely, so there was a lot of freedom in choice. The department head, Jaeger, would appoint somebody with an idea that he might work generally in, say, seismology but what he did was up to him. [chuckle] He had to formulate his own problems.

Listening to you, I reflect on how similar many aspects of my own career were, despite there being a time difference of 20 years or so. But it seems to me that in these last 20 years things have moved much more rapidly than in earlier periods.

Well, life was more leisurely. I think that is a difference in the research environment nowadays – people have to work much harder. I suppose they're all stimulated by each other and have less time outside the lab. At Cambridge we used to go off and play tennis in the afternoon, but here we don't have time for that.

Is the work any better as a result?

The quantity may be greater, but I'm not so sure about the quality.

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Moving firmly into experimental rock deformation

You were appointed to the ANU by Professor John Jaeger, who today has reached almost mythological standing in the Research School of Physical Sciences. Can you tell us something about the circumstances of your appointment to the then Department of Geophysics? It would be unusual today for someone to come into this field from outside the earth sciences community as you did.

It would be somewhat unusual, yes, but it can happen. The background to my coming here goes right back to the beginning of the university and the setting up of the school. It was decided that geophysics would be one of the areas included and a meeting was held in Canberra to discuss where it might go. One of the people brought in to that meeting was Tuzo Wilson, a famous geophysicist from Canada, who suggested that experimental rock deformation would be a good field for the new university to enter.

Oliphant, the first Director of the school, took this up. And when he appointed Jaeger, who arrived at the beginning of '52, he passed this on, together with the thought that it would be worth writing to Orowan for possible names. (Oliphant had known Orowan before the war by giving him a place in Birmingham when he came as a refugee from central Europe.) Orowan was my PhD supervisor in Cambridge, and he put my name into the ring. So I got a letter out of the blue from Jaeger, asking me whether I was interested in this field. And that's how I came here.

No advertisements, no applications?

There may have been an advertisement, I'm not sure.

When you got here, what were your marching orders?

To work in experimental rock deformations – nothing more specific than that.

And starting from scratch, no equipment?

Yes. There was nothing in the laboratory at that time. I first got an X-ray diffraction outfit and then started thinking about deformation rigs. I built two or three rather primitive ones before I finally got onto the sort of apparatus that I use now in the lab.

Was this a new area of apparatus development, or did such rigs exist?

There are two pioneers that one must mention. One is von Karman, whose experiments in about 1908 were quite out of their time. The other is David Griggs, who started in Harvard in about 1935, with inspiration from Bridgman on the experimental side. When I came to the ANU, Griggs had been in UCLA since the late '40s and he had the experimental deformation laboratory in the world at that time. After about four years, in 1957, I had the opportunity to go to Griggs' lab and make contact with him.

So you weren't daunted to enter this new field from outside the area?

Well, from the physics point of view I wasn't [chuckle] because I was really interested in getting into deformation of non-metallic materials. From the technological point of view I was rather daunted; I felt I didn't know anything about high-pressure experimental work and I was a little hesitant.

How long did you take to get your first publications out?

At first I used the piece of apparatus that I brought with me from Melbourne to work on reverse deformation of metals, an aspect of the fatigue of metals work that I had been doing at the Aeronautical Research Laboratories at Fisherman's Bend. I carried on with that here for a couple of years while I was accumulating equipment for the experimental rock deformation work, and I got a publication out on that. But it was two or three years before I got anything out on the rock deformation.

How would you have fared in today's climate of Australian Research Council reporting et cetera?

[laugh] I don't think I could have developed any of the equipment I have now, because I am very slow at things and it takes years to make these things work.

I guess a lot of the work in the School of Physical Sciences was characterised by long-range, very careful instrument development and the science was often very slow in coming. But when it did come, it was invariably outstanding.

And all backed by superb workshop facilities and laboratory assistants.

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A new career in equipment building

Can you tell us something about your time since formal retirement? I say 'formal' because I know that you have not gone fishing but have developed a new career as a builder of scientific instruments.

Well, I had many research students and sabbatical visitors over the years who used the high-pressure equipment that I'd designed and had built in the lab, and they seemed to manage it quite well. It seemed to me that the equipment was reasonably user friendly, and I knew there was very little such equipment about.

I am talking about equipment for deforming rocks at high temperature and high pressure. It is basically a pressure vessel, a big cylinder of steel, in which one generates a high pressure. The pressure medium is argon gas, and when you pump that up to 3000 or 5000 atmospheres it has about the density of water – that is really high pressure for a gas. Inside the pressure vessel you have a furnace which raises the temperature to 1000° or so. In effect, you operate a mechanical testing machine in that environment, applying directed stresses to the piece of rock so that you can plastically deform it. Then you have to measure the forces involved, and we do that inside the pressure vessel so as to avoid problems with friction and pistons.

That took many years of development. I was conscious that although a number of people over the years had set out to build such machines, many had never worked, so I thought that there was room for one that it was viable. So before I actually retired I had the idea that we might go commercial with it.

The Instrom company, a big testing machine company in England, expressed interest in it for a while but finally they said that they liked to make machines at 100 a year [laugh] and they lost interest in my much smaller proposal. I mentioned to a friend of mine in the United States that I was thinking of giving up, but he said, 'Well, why don't you do it yourself?' After some consideration I thought, 'Why not?' and that's how it came about.

So how many have you been making a year?

About one. [chuckle] But we have made 12 so far – they are in England, Germany, Switzerland, France and the United States so far – and we have got feeler expressions of interest from China and Italy. The ones in use are all in earth science departments except the one in Poitiers, France, which is in a material science department. I have just come back from a meeting in Orleans at which the Poitier man gave a talk about the work that they were doing.

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Whither rock deformation studies?

You have witnessed 50 years of change in rock deformation studies, and 50 years of change in the earth sciences. Where are things going now?

Things are already going in the direction of higher pressures and higher temperatures for application to deep material of the Earth – in my view, a pretty limited field. Once you have solved one or two problems there, that may be the end of it. The crust of the Earth has infinite variety in it and I think there is a lot still to be done.

What are the big issues that have to be resolved?

I don't know whether they are big issues but we do need a lot more understanding of how polymineralic rocks deform. So far we have been doing very simple things like marble, which is just an aggregate of calcite; quartzite, which is just quartz; and a dunite which is just olivine. Now we need to do more on rocks which are mixtures of crystals, for example granite, which has quartz and feldspar and micas all in the same rock. We don't really know very much about the deformation of such materials.

Mervyn, you have had a remarkable life and continue to do so. If at any stage you could have changed directions, would you have done anything different?

I'm not sure that I would. I think life consists of taking the opportunities that arise. If I had been given different opportunities I suppose I'd have gone in different directions, perhaps into history or something like that.

I guess there were no jobs for historians, back in the late '30s.

[chuckle] Well, some people survived in that period.

Yes.

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Introduction 

Professor Brian Anderson was born in Sydney, Australia in 1941. He attended the University Sydney graduating with a degree in both engineering and mathematics. Professor Anderson received a PhD from Stanford University in the mid–1960s and stayed on as a faculty member before returning to Australia in 1967. He worked as a Professor of Electrical Engineering at the University of Newcastle until 1982 when he moved to the Australian National University in Canberra to found a new Department of Systems Engineering within the Research School of Physical Sciences and Engineering. In 1994 Professor Anderson oversaw the establishment of the Research School of Information Sciences and Engineering and was the School’s Director until 2002. Between 1998 and 2002, Professor Anderson was President of the Australian Academy of Science. From 2003–06, he was Chief Scientist of the organisation National Information Communication Technology Australia (NICTA). Professor Anderson continues his passion for research in his role as Distinguished Professor at the Australian National University.

Interviewed by Professor Neville Fletcher in 2008.

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© Australian Academy of Science

Introduction

Isobel Bennett is one of Australia's most distinguished and best-known marine biologists. She is unique, in that she achieved this pre-eminence without the benefit of a university degree in her discipline, and in an era when it was very rare for a woman to be a scientist at all. But to counter these drawbacks and to prevail, she was generously endowed with curiosity, tenacity of purpose and high intelligence.

Her earliest work concerned plankton, and she was involved in the first study of it to be undertaken in Australian waters. Another area which she made her own is the intertidal zone of the temperate shores. Her work here spanned nearly a lifetime of meticulous observation.

Her best-known work, though, was concerned with the Great Barrier Reef. As late as 1959 it was said that knowledge of the Reef of a positive kind was far from complete or satisfactory. If this situation has been rectified, it is largely a result of the work of Isobel Bennett.

Interviewed by Ms Nessy Allen in 2000. (Funded by the Australian Government as an ongoing project from the 1999 International Year of Older Person)

© Australian Academy of Science

Dr Elizabeth Dennis is an eminent plant molecular biologist and a Chief Research Scientist working in the Division of Plant Industry of CSIRO in Canberra, where she leads a large team of research workers. Her work has been recognised by many invitations to speak at international meetings, and she is a past president of the Australian Society for Biochemistry and Molecular Biology. She was elected to the Australian Academy of Technological Sciences and Engineering in 1987 and to the Australian Academy of Science in 1995. In 2000, she was joint recipient of the inaugural Prime Minister's Science Prize.

Interviewed by Professor Frank Gibson in 2000.

Contents

• Introduction
• Early years: expectations and opportunities
• University years: nucleic acids and a social conscience
• The native rodents of Papua New Guinea
• Why do plants contain haemoglobin?
• How do plants flower at the right time?
• Gene regulation: plants can't run away from a flood, so why don't they drown?
• Switching on in science: new discoveries, new techniques, new successes
• Putting more iron into the rice bowl
• Beyond the work cycle
• How can we promote discovery-based science?

Introduction

Dr Elizabeth Dennis is an eminent plant molecular biologist and a Chief Research Scientist working in the Division of Plant Industry of CSIRO in Canberra, where she leads a large team of research workers. Her work has been recognised by many invitations to speak at international meetings, and she is a past president of the Australian Society for Biochemistry and Molecular Biology. She was elected to the Australian Academy of Technological Sciences and Engineering in 1987 and to the Australian Academy of Science in 1995.

Early years: expectations and opportunities

Do you think something in your family background, or perhaps your schooling, Liz, led you to choose to study science?

I come from a fairly middle-class family. My father was an engineer; my mother worked on and off while she was looking after the kids. We lived in Hunters Hill, in Sydney – quite a nice suburb, if a little bit ‘bohemian’ in those days (but much more up-market now). A lot of interesting people lived there.

I was the eldest of three girls and my parents were very supportive of any academic aspirations I had. I think my father would have liked me to be an engineer, like him and his father before him, but I always had the expectation of going to university and then following a career. I went to Methodist Ladies College, a girls’ private school which was very good, and unusually supportive of women. Its philosophy was that you shouldn’t not do anything because you’re a woman, and so it provided courses for us like physics honours and chemistry honours, which were unusual then.

As a young girl I was always keen on chemistry. Reading stories of Madame Curie, I decided I wanted to be like her. I think she was the only heroic figure I had in my early childhood. Then at MLC we had a very good chemistry teacher – she had a PhD in chemistry, and was outstanding in those days – who gave us a real interest in chemistry.

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University years: nucleic acids and a social conscience

Would you tell us about your university years? Did you find your undergraduate course – majoring in chemistry and biochemistry – very satisfying and stimulating?

Intellectually, in general no. There were huge numbers of people at Sydney University then : we’d have a lecture of 1000 people and virtually no stimulation by the staff or contact with them, in the first couple of years. I’d say the undergraduate teaching was very poor, except for the ‘odd’ courses. In first year we had Hans Freeman, a very exciting lecturer, for a special chemistry course. And then Harry Messel and Stuart Butler – both exciting – ran a special physics course. In second year, hearing Gerry Wake talking about nucleic acids made me decide that was what I really wanted to do, and later one of the best things that happened to me was having Gerry as an honours supervisor. He taught me a lot about science, about being rigorous, about being focused and methodical.

I went on to do a PhD with Gerry at Sydney in nucleic acids, on how bacteria replicate their DNA. Then I went to New York to Julius Marmur, who was one of the pioneers in physical studies on DNA. He had worked out the relationship between base composition and melting temperature of DNA and also base composition and density in caesium chloride gradients, and developed one of the early methods for preparation of DNA from bacteria. So he was really a bacterial DNA expert, but when I was there he said, ‘Well, what do you want to work on?’ It was very laid back. I looked around and saw they’d started to work on mitochondrial DNA replication in yeast, a natural progression from the bacterial DNA replication work I’d done with Gerry.

During this period you were fairly socially conscious and got involved, I gather.

Yes. The late ’60s was the time of all the civil rights activity and later the anti-Vietnam days. At Albert Einstein College of Medicine, in the Bronx, I found a very active group of doctors, students and post-docs keen on those issues, but also worrying about civil and medical rights for poor people, blacks, and other disadvantaged groups. That whole swag of social issues in New York at that time opened my mind.

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The native rodents of Papua New Guinea

Perhaps because of this social awareness, you didn’t come back to work on DNA, did you?

No, I went to lecture at the University of Papua New Guinea. This may be a paternalistic view, but I thought it was good to do something to use your knowledge for the less developed countries. That was quite a mind expanding experience. I think normally we are comfortable in our own little niches, but Papua New Guinea didn’t have all the molecular biology, high-tech instrumentation and big infrastructure, or an environment where you can feed ideas off lots of colleagues.

I was doing chromosome and DNA studies on the native rodents, and together with a classical zoologist, Jim Menzies, I would go out trapping rodents and taking pictures of them and determining the chromosome complement. We even wrote a book on the rodents of Papua New Guinea. There is a big radiation of species. Some of them are cute little tree mice but one rat is called  Hyomys goliath because it’s so big (about 400mm head and body and tail the same weighing 1kg). And they’re quite unrelated to  Rattus. That project was fun and I saw a lot of the PNG countryside. Doing this work taught me that there are a lot of scientific problems and it’s interesting when you get involved in any of them.

Why do plants contain haemoglobin?

Later you moved to the CSIRO Division of Plant Industry, in Canberra. Were you pleased that that meant you were able to get back to your beloved DNA and to develop a long-term research program?

Yes. At first, I worked with a very good group under Jim Peacock’s leadership, including Doug Brutlag and Rudi Appels. We looked at repeated DNA in  Drosophila. Then I returned to Papua New Guinea for a while. When I got back to CSIRO, in 1976, the mission of Plant Industry had become more focused on plants, so we switched over – still in conjunction with Jim – to working in DNA from plants.

During the ensuing years you’ve covered several topics. Could you tell us something about the work on haemoglobin? That doesn’t sound very plant-like.

No, it doesn’t. That was quite an exciting development. It was inspired by Cyril Appleby, who is also a member of the Australian Academy of Science. He worked on haemoglobins, initially from legumes which use haemoglobins to carry oxygen past the bacteria involved in symbiotic nitrogen fixation. Then he isolated and worked on a haemoglobin from an Australian tree called  Parasponia, which also fixes nitrogen. We decided we would clone the Hb gene  Parasponia. It was known, as the basic principle, that haemoglobin is important in fixing nitrogen in the nodules of legumes and that other Australian trees also fix nitrogen. The question was whether the gene that is used in the non-legumes is the same as the gene used in the legumes. So we isolated genes from other species – in the beginning, from plants in Australia and Papua New Guinea like  Casuarina and  Parasponia, which fix nitrogen.

We found that in fact there were two quite different Hb genes, one that was in the legumes and also in  Casuarina, and another one in  Parasponia that was quite different. But we then found that the gene in   Parasponia  was also present as a second gene in  Casuarina, but not in the nodules. It became clear that there were two families of Hb genes, each of which had been recruited to fix nitrogen, one in the legumes and one in the  Parasponia. We then went on to show that in  Arabidopsis, which has nothing to do with nitrogen fixation, there are two classes (and now possibly a third) of plant haemoglobins. We have extended this to suggest that Hbs are in all plants. They’re related to the animal haemoglobin, so they’ve probably been there ever since the divergence of animals and plants about 1500 million years ago. Hbs must have had some function other than nitrogen fixation in plants, and then subsequently, as the legumes and  Casuarina and other nitrogen fixing plants evolved, the haemoglobin genes that were there became adapted to function in nitrogen fixation. If you cut open a nodule on the roots of a pea, you find it’s quite red because it’s got haemoglobin in it.

We’ve been trying to work out the function of these Hb genes in plants and the only thing we’ve found so far is that one of these genes in  Arabidopsis is switched on by low oxygen. In fact, if you put more of that Hb into the plant, it protects the plant against low oxygen stress. So that may be the function of one, but we have no idea what the function of the other one is.

How long have you been working on that problem?

For 15 years, on and off, mainly working with students. As new techniques become available, we go back and try again. Of course, all the work is a lot of collaboration with different students and postdoctoral fellows.

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How do plants flower at the right time?

You also have an interest in the flowering of plants. In general terms, what is that about?

That is very much our current project, involving all the people in the lab. Flowering is an important developmental process in plants, a very exciting biological process. In animals you set aside your germ cells early in development, but in plants there is a growing point which starts by making leaves and stems and vegetative structures but then switches to making reproductive structures like flowers and pollen, all the components that make up flowers. So how does that switch occur? How do cells derived from the one growing point change from making vegetative to reproductive cells?

Using  Arabidopsis, we isolated a mutant that didn’t flower till very late, and then we isolated the gene that was mutated to cease this effect. We could show that this gene acts as a repressor of flowering: the more of the gene product there is – the more the gene is switched on – the later the plant flowers. It’s a quantitative controller of flowering time. Later we found that this gene is down-regulated by vernalisation, a response to a period of cold which has long been known to cause plants to flower.

Plants need to flower in the springtime, not when there might be frosts or when they won’t have enough time to get their seeds mature for the next generation. They can’t go inside when it’s cold, so it is very important to them to flower at the right time and they’ve evolved mechanisms to ensure this happens. One of those mechanisms uses the cold as a signal. That is, many plants require a period of cold – a cold winter – in order to flower in the spring. They may also use day length (they recognise when the days get longer) as well as vernalisation to ensure that flowering occurs at the right time. We’ve found that the vernalisation, or cold, switches off the repressor gene called FLC – flowering  LOCUS C. After this repressor of flowering is switched off by the period of cold, the plants flower. So we have a molecular basis for one of the long-standing question in plant biology – how does vernalisation work?

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Gene regulation: plants can’t run away from a flood, so why don’t they drown?

You’ve also been interested in the regulation of plant genes, haven’t you?

Yes. That’s the basic theme that makes it all hang together. The major project we’ve had, over a long time, is looking at genes being switched on by low oxygen, such as occurs in flooding or waterlogging. When plants are flooded they can’t run away. If it gets wet for you and me, we just walk away, but the plants are stuck. Plants have evolved mechanisms to cope with stresses caused by their being sedentary. So, when plants are flooded, they switch their metabolism from oxidative metabolism – oxidative phosphorylation – to fermentation. They make alcohol: the genes for the ethanol fermentation pathway are switched on. Instead of pyruvate going into the Krebs cycle, it enters the ethanol fermentation pathway where it is converted first to acetaldehyde and then to alcohol, using the enzmes pyruvate decarboxylase and alcohol dehydrogenase.

Over the years we have looked at these genes switched on by low oxygen, trying to identify the promoter motif so important for switching them on. In fact, all these genes have the same anaerobic response elements, so they all have a DNA sequence, upstream of the gene – a promoter element – that’s important in the glycolysis and low oxygen metabolism alcohol fermentation pathways. We’ve now identified the protein that binds to that low oxygen control element. That’s the sort of thing we’ve been interested in, with a view to trying to help Australian agriculture by making plants more resistant to waterlogging.

You had an interesting time at Stanford, I understand, working on gene regulation.

Yes. In 1982–83 I spent a year’s sabbatical as a Fulbright Fellow in Paul Berg’s laboratory. That helped me in several ways. For one thing, it helped me to understand the thinking that people in Paul’s lab and other labs in Stanford were using to try and analyse the control sequences of genes. They were further advanced toward understanding the sequences that replicate genes. To introduce a candidate you introduce it back into animal cells, see what effect that has, irradiate mutate a few bases, and see if this alters expression of the gene.

But also, the whole way the Stanford biochemistry department worked was very good for me in considering how laboratories, departments, could work. That department was very distinguished: two Nobel prize winners were in it – Arthur Kornberg and Paul Berg – and maybe three-quarters of the rest of the faculty were in the National Academy and really were very innovative and exciting scientists, like Dave Hogness, Ron Davis and Dale Kaiser. Charlie Yanofsky, in biology, was around there too. The department was small, with only about eight or ten members of staff, but the way they worked together produced a very exciting intellectual environment. They had staff meetings where they discussed science and the department, they pooled their grants, they worked together, they didn’t try and look after their own equipment or prevent other people using them. Everything was cooperative, which seems a very good way to work.

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Switching on in science: new discoveries, new techniques, new successes

Over the years, in your various activities, you have made quite a few novel discoveries. As a result, you have been invited to talk at many international meetings.

Well, it’s been the right time, with the new techniques in  Arabidopsis and new expression techniques in plants, for us to identify what the sequence is controlling the genes switched on by low oxygen. The completely unsuspected idea that there are haemoglobins in all plants is also a novel discovery, and then this gene that regulates flowering, that we’ve recently been working on, is pretty novel too.

A lot of this work has been done in collaboration with other people, and so we share around the talking at international meetings. And I’ve had a long-term working arrangement over these years with Jim Peacock, who has had a big input into projects.

Would you say there have been major switches of your research direction at particular times? If so, what caused them? Which gene did you switch on?

I think changes in direction come with the opportunities provided by new techniques. The advent of cloning, of being able to isolate genes, has certainly been important. Initially we worked on repeated DNA in  Drosophila and in plants. The  Drosophila DNA and also one of the plant DNAs have the sequence like AAGAG, and it took us about 3 years to sequence those five bases! This little five bases repeat was repeated maybe a hundred thousand or a million times, but to actually get that sequence – back in the middle 1970s, with Doug Brutlag and other people – took a long time. But then the ability to sequence DNA, and later, cloning, enabled us not just to look at repeated DNA, but to isolate the genes and to study them in detail. The technology has been tremendously important. The development of  Arabidopsis as a model plant, with its genome now almost completely sequenced, has been very important in plant molecular biology. So often, it seems, we spend years doing something that appears really simple afterwards!

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Putting more iron into the rice bowl

You have also applied your interest in biotechnology to rice, in a very important project. Has the outcome been satisfying to you?

This project built on our work on haemoglobin, a protein that has a haem group, which binds iron.

People in developing countries who eat rice as their major food source face two major problems of nutrition. One is that people can go blind from vitamin A deficiency, and the second major problem is anaemia. Rice doesn’t have much iron, so something like a billion people suffer from iron deficiency anaemia. One of our projects, with support from CSIRO and from the rice growers of Australia, has been to try and increase the iron content in rice. We’ve added genes for haemoglobin under the control of strong regulatory sequences so that they’ll have high levels of activity in the seed and the seed will contain large amounts of haemoglobin.

So we’ve got transgenic plants with high levels of haemoglobin, but we don’t really know how much that has done for the iron, because it’s very important how iron is presented when you eat it. If it is associated with haem, it’s much more bio-available and can be absorbed: it can be 20 times more efficient than if you just eat the same amount of iron as iron filings or ferrous sulphate. So we’re starting feeding trials on rats – and there’s an artificial system now – to see whether we have improved the nutritional properties of the rice as far as bio-available iron genes.

The problem of the vitamin A has been attacked by Ingo Potrykus and other workers in other parts of the world, and you may have heard they have had some success in making rice that contains higher levels of vitamin A.

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Beyond the work cycle

Apart from being President of the Australian Society for Biochemistry and Molecular Biology, for a long period you’ve been on numerous government committees and so forth. Do you regard your committee work as an act of charity or a necessary ill, or do you really enjoy it?

I guess a combination of all that. There is a duty side to it: I think we have a responsibility to put back something into the scientific community so we should do some of this, just to help. But I particularly enjoy committees which are reviewing science and trying to see what people are doing, trying to evaluate it and make suggestions for where things should go. I have enjoyed being on committees reviewing departments, or research grants committees, because I like broadening my focus. I learn from it too. That side of it I like; some other committees are more duty.

In such a very busy life, I imagine you’ve worked very hard and constantly. Do you have any outside interests at all? If so, how do you fit them in?

Well, I have a family – I guess that’s an outside interest. We have two boys, now aged 15 and 13 so they’re pretty active, and looking after them takes a bit of work. They’re probably not looked after as well as they might be, but they seem to be pretty tolerant about having a mother that’s a scientist, and the idea of my having to work. Most nights, I come home, make dinner, have a bit of family life and then go back to work about 9 or 9.30 and work till about 11 or 11.30.

My other major interest is that for just on 20 years a group of us have owned some land on the coast, at Tilba. It was a dairy farm, so it had been completely cleared – there was only one tree left on the whole 70 acres. We do a lot of tree planting and revegetation, and have enjoyed seeing what happens when you restore the vegetation, watching what birds and animals come back. It’s right on the beach, so it’s also a very nice place to be: it is very relaxing to enjoy the botany and wildlife and have a swim. Next to us a national park has been declared, so we’ve been active in getting it protected, stopping people from driving on its dunes, and revegetating those dunes too. That’s been very important, and to have that break away from work does help – no telephone, no electricity down there.

How can we promote discovery-based science?

You have some pretty firm theories on what should be taught at university, and how.

I really enjoyed university, but largely because I lived in Women’s College and so I made quite a lot of close friends. We’d spend time discussing the meaning of life and the intellectual basis of ideas. In the days when I went through university, it didn’t develop people’s capacities to their full or take full advantage of their abilities. I would have preferred smaller groups doing projects, where people actually focused on things that they needed to know – trying to work from a project-oriented base rather than having a vast amount of knowledge that had to be just stuck into you. Kids now are not reading anywhere near as much as they used to, but they are very computer literate and they do learn in a much more empirical way than we used to. I think that education could change quite a lot towards taking advantage of the flexibility of younger people and their ability to pick up things as they need to know them. If you can give them a project where they need to know things, and they go out and find them, that is a much better way to learn.

I learnt a great amount of organic chemistry that I’ve completely forgotten because I’ve never needed it again. I think that very classical view of education we’ve had should be changing, but I can’t see many of the universities responding to that.

There has now been another change in biology, into genomics: instead of looking at single genes, we have to look at whole genomes and see all the genes that are changing and what’s going on. That’s much more information-rich and information technology-rich science and again has a much more empirical basis. You might start off with a hypothesis, but part of the question will be, ‘What gene is switched on?’ rather than, ‘Let’s test if gene X is switched on.’ So the science is discovery-based rather than hypothesis-based. You might find a lot of new parameters that you didn’t even think about. If you start off in the traditional way you say, ‘The genomics approach means that you’ll find X, Y, Z and others that you’ve never even thought of’. It is a much broader view of science than we’ve been used to.

You talk about the genomics approach. Presumably you’re thinking now of actual research work rather than instruction?

Yes, that’s right. But if we’re instructing people so that they can use the new science, we have to encompass this new approach to research in the teaching as well. There are two pressures for that. One is the sort of research that’s being done, and the second is the sort of background the kids coming to university have now, which I think is very different from what we had. We were all reading a good deal but had a narrow base of experience. But I think kids come now with a much broader background – not as intent on reading, much broader because of the computer and the internet. And so instruction has to change too. The way to link those two things is to give them at least project-based work: not ‘Here’s a mass of knowledge, learn it,’ but, ‘Think about doing this, and find all the things that need to come out of it.’ There’s so much knowledge, you can’t hope to learn it all. How do we decide whether we should have five hours a week organic chemistry, or one hour a week? What’s the basis? What is the body of knowledge that should be passed on? We don’t know.

Thank you, Liz, for giving us so much insight into your experience and your ideas.

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Professor Jonathan Stone was born in 1942 in Auckland, New Zealand and moved to Australia with his parents when he was a baby. He received a Bachelor of Medical Science from the University of Sydney in 1963 and a PhD in 1966. After a year as a visiting lecturer at the Hebrew University of Jerusalem, Stone did post-doctoral research at the Institute of Biomedical Research in Chicago with Sir John Eccles, and then spent a year in Munich at the Max Planck Institute for Psychiatry. Stone returned to Australia in 1970 to take up a position as a research fellow at the John Curtin School of Medical Research. Here he continued his research on the retina. In 1976 Stone moved to the School of Anatomy at the University of New South Wales (UNSW) initially as a senior lecturer, then as an associate professor (1978–85). He became head of the school and took up a Personal Chair in Anatomy in 1985. During his years at the UNSW, Stone's research interests shifted from the study of parallel processing to the development of the brain. His next appointment was as Challis Professor of Anatomy at the University of Sydney (1987–2003). Here he worked on the interaction of neuroglial cells during the stresses of birth, particularly focusing on types of blindness that result from the degeneration of photoreceptor cells. Stone was appointed Director of the Research School of Biological Sciences at the Australian National University (ANU) in 2003. At the ANU his research concerns the stability and degeneration of the central nervous system, including dementia and a group of inherited eye diseases that affect the retina.

Interviewed by Dr Max Blythe in 1996.

Contents

• Cradled with warmth, intellect and no nonsense
• Knowledge, philosophy and cheerfulness
• Retinal insights: developing the whole mount
• From the retina to receptive fields
• Working with Jack Eccles
• A parallel-processing odyssey: working again with Peter Bishop
• The family man and sportsman
• Moving on: balancing teaching with brain research
• Building up research
• Blindness: the ontogenetic link
• A wider arena: Alzheimer's disease
• And wider still: animal welfare
• Population biology
• Optimist or pessimist?

Cradled with warmth, intellect and no nonsense

Perhaps we could start with your family – the cradle of your attitudes and achievements.

I was born in Auckland in 1942, three years after my parents had gone there for my father to take a Chair of Law. (He was very young, in his early 30s.) As a babe in arms I was brought to Sydney, where my father had been appointed to the Challis Chair of International Law and Jurisprudence. My mother tells me we came in one of the old QANTAS flying-boats and landed out on Rose Bay.

As you grew up, the family environment that moulded you was fairly tough.

Yes, it was. My parents were by then immigrants to Australia but their own parents had been immigrants to England from Russia, fleeing the Tsar's armies and their claims on the Jewish population in Lithuania. They were fighting their way up to the middle class, and doing a good job of it. From the slums of Leeds, my father went to Oxford just at the time when it was opening up. Although he recognised that it gave him his start, and on his study wall he kept his college shield – Exeter, I think it was – he had a love/hate relationship with the place. He was never invited back despite all the distinctions he gained, and didn't himself ever want to go back, yet he spoke with great affection of his tutor there, Geoffrey Cheshire – Leonard Cheshire's father. He kept up contact with the elder Cheshire for many, many years.

They were tough people who had come through tough times, a very strongly Jewish family. That was the time when the British mandatory powers in Palestine were being debated, and as a young man, newly arrived in Australia, my father took a very strong stance against our Governor-General. When I read those old papers I can sense the mixture of intellect and anger that was driving him. But that was not part of my life. Being just a babe, I was unconscious of it. Then I found myself in postwar Australia, growing up, going to school.

I think your father was a strong, powerful loner, perhaps not unlike you.

Yes. You never know whether you are imitating somebody or you just are that way yourself, but my instinct has been to follow my mind. I have never had any problem with authority of the institutional sort but I have always had a problem with intellectual authority.

You got into a few conflicts.

Yes, which with a bit more of a smile and a shrug and a tug of the forelock I could have just gotten round, but my instincts weren't there. They are well in place now, with all the learned reflexes, but I do not think I somehow or other inherited them.

I'm sorry I am never going to meet your fascinating father. Tell me about the rest of the family: your mother, brother and sister.

It was a close marriage. My mother, Rebecca – Reca, as we call her – is still alive. She devoted herself to the family and provided the traditional background which my father drew on. My elder brother has for many years been Professor of Religious History at the Hebrew University of Jerusalem, and my sister is a psychiatrist practising in Sydney. They have very stable marriages and happy families, and ours is a very warm sibling relationship.

Mother was strong, as well as Father?

Yes. She came from that generation that had fled and then lived. Luckily my parents found themselves in England and therefore unaffected by the Holocaust, but they were fiercely burnt by it psychologically. That gave a toughness, a no-nonsense attitude. For example, I didn't do much sport at school because it just wasn't encouraged – what mattered was the knowledge, learning, getting qualifications. I have since made up for that in various ways, however.

I became aware of the Holocaust only in the most peripheral way, although of course as a kid you learnt about it. But I realised that within central Europe something of the most awful nature had happened. Out of that I drew a scepticism about the role of culture – Kultur, in the German sense – and I realised the amorality of culture. Like my father, who at core was intellectual, a common lawyer, I came to appreciate the value of the British tradition: a powerful set of mechanisms, quaint and odd and full of idiosyncrasy, which nevertheless had provided England and could provide Australia with a form of government that, for all its imperfections, got rid of that terrible instability that afflicted middle Europe. And I thought about that, trying to understand it at its fundamental levels.

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Knowledge, philosophy and cheerfulness

Did your primary schooling have much influence on you?

No. After the local school, Gordon Public, which I remember with affection, I spent two happy years at one of the selective primary schools which were called 'opportunity schools'. Then I passed into a selective high school, North Sydney Boys. That was a tough environment, not really a caring one – not care-less, though – and I have since learnt through my children's experience how much more caring and attentive schools can be. But it was a fair environment, which I appreciated. I threw myself at the academic windmills there and did well enough. I wanted to have a go at everything, but the area that I really found myself standing out in was English: for some reason the words came and I could be a star at that subject.

I suspect it was through parental pressure that you went into a medical course at Sydney.

Yes. I came out of high school young, only 16, with no clear idea of what I wanted to do. Hell, I thought I was ready for anything. I'm sure my parents would have supported me in doing any of a number of professions, but I wasn't sure which I wanted. They suggested medicine, and in I went.

Going in at 16 couldn't happen now, could it?

No. School has been extended by a year, for a start, and there are increasing attempts to bring people into medicine at an older age. And I was a case in point. I threw myself at the medical course. It was then, even as it is now, a flood of knowledge that people package for you as best they can. It didn't help that those were extraordinary years: after the war, the Australian universities were asked to cope not just with the normal flow of kids coming out of high school but also with the returned servicemen. They were not allowed to put academic restraints on. There were 700 of us in that year, of whom 300 failed, and when we passed on we picked up 300 more that had failed. It went on for a few years like that, during which we were dealt with en masse. I wasn't enjoying it, and when the opportunity came to do a research degree, the Bachelor of Medical Science I quit the medical course and went into research.

You liked writing and I've got a feeling that you were reaching out for quite a bit of philosophical material. Was that linked with your father?

Yes. I got from him, whether through genes or absorption, a need to understand at some fundamental level what was going, without getting into either the scholasticism of a lot of modern philosophy – which I read and just couldn't handle and then came to understand for what I think it was – or the gloom that Boswell referred to in his Life of Johnson. (He commented that he had tried to be a philosopher but cheerfulness kept breaking in.) At some point you walk away from it, because it just drags you into deep and difficult debates, but you realise that the philosophers have tackled very difficult issues that afflict us generation after generation, and I found I could make my way better in science if I understood those issues.

You were already strongly analytical. Did science help to further that?

Yes. Science gave a lot of materials for understanding. For example, I found it fascinating to read about the debate over evolution – not so much the debate itself, which was a confrontation between two systems of knowledge, but the drive in some people within the scientific system to have a clear-cut classification that everybody could understand, while others said, 'Wait a minute. What we need is a classification that can deal with errors, that can be creative.' Heuristic is the word that the philosophers use. As I came to understand that, I could see what was drawing my colleagues in certain directions. I felt I became a fuller scientist for doing that.

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Retinal insights: developing the whole mount

Perhaps you can put me in tune with the environment that Peter Bishop had created for that Bachelor of Medical Science degree.

Peter had the passion in him to establish the study of the brain in Australia. Eccles had a group in Canberra, in the very specialised environment of the Institute of Advanced Studies, but there was almost nothing else. After Peter had been to University College in the late '40s, he set up major labs at the University of Sydney, he started to bring in people and I was one of those lucky enough to go into his lab. Some were extraordinary people, who have gone on to make their mark right throughout Australian neuroscience and overseas. And in Canberra he attracted a lot from overseas who later made their mark in their own countries. I feel privileged to have been one of those people.

Peter was interested then in the cerebral cortex. He had moved on from the study of individual neurones, which he did so well, to studying the system as a whole, going to the highest levels. The first project he gave me was to explore the visual cortex but I couldn't do it. He was so busy that I found myself on my own, and I remember gritting my teeth and saying, 'I'm just going to work till I find my way out of this.' The way out came partly through a collaboration with Bob Rodieck, an American – extremely bright, full of ideas, full of self-confidence – who now has just about retired after a very distinguished career. I was lucky to meet him at that stage. The group included also Bill Levick, who has just retired from a personal Chair at the ANU, and Jack Pettigrew, who now heads a research centre at the University of Queensland.

But also I created something myself, when I got really interested in the structure of the retina. Peter gave me an experimental project on the retina, the starting point of the visual system, and I had an insight into how to look differently at the retina. I drew on old sources to figure out a way of taking the whole retina and laying it flat, instead of cutting it up into pieces. This proved to be an enormously powerful way of surveying it. I presented that at my first scientific meeting, of the Australian Physiological and Pharmacological Society in Melbourne, in 1962. I can remember during the long train ride back from Melbourne after that meeting thinking how to use the technique to explore new features of the retina.

Is the retina easy to peel off? Does it dislodge easily?

Once you get confidence, yes, and now the method is used in hundreds of papers in many, many labs every year. Although its origins are forgotten, I have some pride in having contributed at that point. I must have been about 19 or 20 when I made those first whole mounts. And I'm still looking at them – I spent time on the microscope on one this morning. It's a powerful technique. Of course we have brought in the more molecular techniques in subsequent years and the new forms of microscopy (fluorescent, confocal), but that was a core thing for me.

The use of whole mounts gave me incisive data, new ways of looking at things, and I was ready to go. We came up with a series of studies then which led to an understanding of aspects of the retina on which Peter could draw to strengthen his cortical work. I've always been grateful to him because, once he understood what I was getting at, he gave me freedom and support to go for it.

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From the retina to receptive fields

Peter Bishop had already looked at the cortex – a mosaic of cell areas, interpreting the messages from the retina – looking at the optic nerve and the geniculate nucleus on the way through. Were you actually providing a fresh look at that mosaic?

Yes. I went out to the periphery.

So you were mapping this retinal structure?

That's right. With all my analytical and holistic desire to see the large picture, I found myself getting down to very detailed counting – one, two, three, four, five – but I did get the data on which I could see patterns of specialisation of the retina. Two of Peter's collaborators (Wladimir Kozak and George Vakkur) had started to look at that but hadn't concentrated on it. I picked up from them, getting down to the microscopic level. That enabled us to see how the retina divided its projection into the two halves of the brain and we began to pick up the morphological differences among the ganglion cells, which then became the basis of the parallel processing story.

After that first period in Peter's laboratory, in the '60s, you did a PhD with him.

Yes. Peter was my supervisor. The study had physiological parts which I did with Bob Rodieck, the American collaborator, and it had anatomical parts which were much more my own. We were working at a point in the study of visual pathways at which people had started to look at the receptive fields of the cells. Once it was realised that the cell would not respond to the whole visual field but only to part of it, you could define the part (the 'receptive field'), analyse it, start to model it, and then when you surveyed many you could start to see the differences. That had been begun in America in the late 1930s by Hartline, who got the Nobel Prize for his work. The work had stopped during World War II but this was its renaissance, certainly in Australia. That work culminated in a mathematical model that Bob Rodieck produced, calling it the convolution model. I went a fair way down that stretch with him – he did the mathematics – and that became a paper of reference in the literature from which then we could begin the classification work.

Peter was quite insistent that for my post-doc years I should go overseas, and he arranged for me to work with Jack Eccles, who by then was in Chicago.

Leaving Peter Bishop's unit was not totally without some stresses with colleagues, because you were burning rather a bright light at that stage.

Certainly with Peter it was a friendly leaving and later he welcomed me back to join the department he then headed at the ANU. With my colleagues, I suppose there must have been something in the chemistry of personalities but the substantive issues on which I found myself at tension with them were conceptual. Although I felt I did do my time painting the fine strokes, perhaps I was more willing than some other people to draw broad strokes.

I knew I had so much to learn from those colleagues, because they had skills and enthusiasms and abilities that I didn't have. But science is a matter of choices. I was always drawn to where I thought we were breaking through into new ground. Creative science is always done on the edge of the unknown. You are always stumbling in the dark. You will be clumsy, fall, look silly, but I was never afraid of that. Others were more content to stay in the safe ground which was a bit explored, where they could do much more elegant work. I rejoiced in what I believed was new, despite my clumsiness. They rejoiced in the (very real) elegance of their work. That was the tension.

So you really were ready to go to Eccles in Chicago in 1965?

Oh yes, I was ready to go.

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Working with Jack Eccles

My post-doc years were exciting. Working with Eccles was a terrific experience. I did intracellular recording. Two of my collaborators were Henri Korn, who is now a major figure in French neuroscience, and Nakikira Tsukahara, who was rising in Japanese neuroscience when he died tragically in an airline crash.

Can you give me a portrait of Jack Eccles himself?

He was a gorilla of a man. He was quite big, still full of energy – in his youth, I think, he'd been a great swimmer. He was positive and quick to judge, and if you were on the line that was his at the time, he was there and supportive. But if you differed too much, dug in your heels, he would just walk away from you. He was in his late prime by then: he had achieved great things and had great self-esteem. I was prepared to absorb, and I learnt a great deal from being in that group. Later, I drew a lot from Eccles' philosophical writings – from the fact that this man of empirical commitment believed he had something to learn from philosophy.

And, as you were saying of yourself, he was comfortable with being on the frontier of science, with getting things wrong and stumbling.

Yes, he was. At one conference I remember, when somebody put forward what was then a fairly radical idea – I think it was the proposition that all the climbing fibre inputs to the cerebellum come from the olivary nuclei – only Jack Eccles would speak up and support it. He just wasn't afraid. He was willing to take a strong position, and he had real judgment. Jack Coombs, who had worked with him, remembered this judgment in one of Eccles' great discoveries. While they were working on a quite different issue, recording synaptic potentials in the spinal cord, they kept seeing something unexpected and irrelevant: a synaptic potential inverted on the screen as the neurone became injured. Jack Coombs told me that after that happened two or three times, Eccles said, 'Forget about what we're doing. That's what we're investigating.' Out of seeing that epiphenomenon came his analysis of the mechanism of this inversion, and later his book on the ionic basis of synaptic transmission, which was his key contribution.

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A parallel-processing odyssey: working again with Peter Bishop

When Eccles left Chicago, you went to one of his first-rank people.

I was offered a position in another lab in Buffalo, where Eccles went, but I'd dragged my family round enough. I stayed with Rudolfo Llinas' group and worked with John Freeman, a young American who showed me how brilliance and good hands could take you a long way very fast. I greatly enjoyed that year with John, going into intracellular recording using classical electrical stimulation techniques, which I then took back to Canberra.

Via Munich.

Yes. I had a year in Munich, partly to see Germany but also because I was lucky to get a position there in the Max Planck Institute for Psychiatry. The head of the lab was Otto Creutzfeldt, a very distinguished neuroscientist whose father's name is attached to the Creutzfeldt-Jakob disease. Otto later headed a group in Göttingen, and Bert Sakmann and Erwin Neher, who got the Nobel Prize a few years ago, were his protégés. He too supported me and left me to my own devices while I brought together the techniques I had learnt with Eccles with the work I had done with Bishop.

From Eccles I had learnt to watch for the unexpected in what you see down the microscope or on the oscilloscope screen. And that's what happened in the transition from Germany back to Peter Bishop's group. When I worked in Germany on a very detailed study of conduction velocity of cells with different receptive field types, I saw variations that no-one had described. Hanging out on the end of the traces were cells that didn't fit into the two major types. I put them aside…

And that's where you stumbled across the W-cells?

Yes. Briefly, when you stimulate along the visual pathway – as Bishop had shown 20 years before – you get two groups of fibres turning up at very precise moments on the oscilloscope screen. When I started to work on their receptive field properties, I saw a very occasional scattering of cells which I guessed to be small neurones that others hadn't seen in the past 20 years because of microelectrode technology. I went back in, using the glass electrode technology that I'd learnt with Eccles, and then they started to turn up, one in five. When I did look at the receptive fields they were quite different, and I realised that I was dealing with a class of cells that hadn't been recognised before.

Were these patterns and populations of cells with very precise influences that turn them on or switch them off?

That's right. And that was the excitement that came out of Hartline's work. Buried small in my doctoral thesis had been a chapter in which I gave – when I look back at it – an incomplete description of receptive fields that hadn't ever been seen before. It was very exciting at the time and it got published in Science, but they were not easy to record and I couldn't do more with it. It stayed somewhere in my memory traces, however, and when I reached back to it I found that although it was a bit untidy it looked better with passing years.

I had five fruitful years with Peter's group. He put me in a lab, with tremendous support. I got technical support equipment; I didn't have to write a grant, just go. The first story that we pulled out was this new class of cell in the retina. Having absorbed that, then I turned to how the retina sent information to the brain. The ganglion cells are the output cells of the retina, and in your eye and mine there are a million of them, each has its own private line into the rest of the brain. Among the ganglion cells there are clearly different cells doing different things. The retina is not just a light processor but is specialised, through these different output cells, to pull different things – movement, shape, colour in the primates – out of the visual image and channel them into different parts of the brain. The idea driving me was that when you then traced them in the brain you would find they went to different areas, and this worked.

I mustn't fail to acknowledge the superb work done by the others working in the same area in Canberra. We drove the idea right through to the visual cortex. With colleagues who made tremendous input to it – Peter Hoffmann, Bogdan Dreher – we argued that these ganglion cell classes are reaching the cortex without mixing and are driving some of the variation in the cortex which American workers had interpreted as serial processing. They had seen the same variation but had thought the information was being processed in a hierarchical manner. One of the papers we wrote together was on hierarchical and parallel visual processing. It contained large areas of analysis that I still am pleased to read.

That driving idea of parallel processing was resisted by the Americans. They had done such a lovely constructive job; they broke open so much, and were rightly rewarded with great honours. But on this…

You were not yet 30 and you were pushing against big, established ideas.

Yes, and was stupidly fearless about the matter. But I must say that Peter Bishop understood it: he saw what was happening and his philosophy was that out of the ferment, which he welcomed, would come the hard testing, the discarding of what was wrong, and so he gave us great intellectual freedom. I like to think that he found that a very productive period in his lab. I believe those ideas are now part of the mainstream. My father used to say in his own field, 'There comes a stage, son, where not only don't they remember the details of the battles, they forget that there ever was a battle.' I think we've got to that stage now with the parallel processing concept of visual processing.

I haven't done the cortical work since then, but we have seen evidence that different areas of visual cortex are handling motion information, information about movement, information about colour. Even the recognition of faces – you would be aware of the work of Oliver Sacks – seems to be packaged into parts of the cortex. I didn't believe it would go as far as it has. Quite extraordinary stuff.

You were drawing now on the Eccles time, those five years out of the Bishop world, as a philosophical time when you probably read Popper – an exciting time of transition.

Yes, although I must admit I didn't really tackle the philosophy until, in Peter's group at Canberra, I started carving out new paths. That's when the fire really started and I found myself under a philosophical attack as well as an empirical one. And that's when I went back to it.

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The family man and sportsman

Jonathan, before we move you on from Canberra, could we look at how you took a wife and became a family man.

I met Margaret as a lab technician in Peter Bishop's lab. She was then studying Arts part-time. We rubbed along well enough in the lab but struck up a relationship after a few years of being together there, and ours has been a warm and successful marriage that I've relied on. I've drawn very deeply on her intellect as well as her warmth as a person. We have three daughters, but after we returned to Canberra she enrolled in law, did wonderfully well, got a job when I moved up to Sydney in 1976 and began her own career. She stayed with the New South Wales University Law School for perhaps 15 years, and is now going strong with one of the big law firms in town. That's been a very positive, warm, stable family life.

But not so strongly Jewish as your background. I think you put aside some of the religious pressures.

Yes. The issue of ethnic involvement is such a fascinating one for anyone brought up in that tradition, because obviously there is a religious core to it. And on issues of faith I am a sceptic, an agnostic. There is a ritualistic core to Judaism, and I am not an admirer of ritual. It has both strengthening and divisive aspects to it. But nevertheless it wasn't a sanctuary into which I could escape, nor did I really want to escape. In fact I have close colleagues in Israel and work there from time to time, and I enjoy and am fascinated by the extraordinary history of that place and time and of those people, of whom I am certainly one.

Are you ethnically associated now, rather than religiously?

That's right. Every religion goes through its own evolution. It seems to me that the evolution of Judaism was stunted by its diaspora, by its being spread out – that now, having its own space and patch in the sunlight, it is starting to evolve again. But that's another story. I am part of that, but just a part.

As your career took off, you began to find the odd leisure opportunity. You become a sailor. You haven't been a sportsman before but have just ground away at it all.

Yes. I had played a few games – a bit of squash and so on. But my enthusiasm, to which I didn't come till after I was 40, is racing dinghies. I realised I had grown up in Sydney, with that superb harbour, and had not sailed. I have since learnt to sail them and to enjoy the racing, and I have as many friends in my small harbourside club as I have in the rest of my career. That has been a wonderful side to the sport.

What did you and your family find Canberra like? It must have been a culture shock.

Yes, but we were at the stage, with young kids, where Canberra was ideal – just laid on for young families. You can complain of Canberra even now that it is less culturally rich than great cities, and that is certainly true. Canberra was and is still a middle-class town. If you take it as it is, it is just great. You can grumble but we didn't. We had a wonderful five years there.

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Moving on: balancing teaching with brain research

You mentioned that you moved next to Sydney. To a senior lectureship, wasn't it?

Yes. I applied for and was appointed to a senior lectureship in the School of Anatomy at the University of New South Wales. Despite the scientific ferment in Peter's department, I found the institute life, the life of turning up to work with a small group day after day, quite stunting, I wanted to be interactive. I was looking for lots of faces. I wanted to teach, very actively.

You had been coming to Sydney to teach, hadn't you?

Yes. I had come up year after year to give a few lectures, and always enjoyed doing it. Also, my father loved to teach and he taught me a lot about teaching, just by example. He became a media figure in Australia. I can still remember him giving his last lecture (he was very sick at the time but he was determined to do it) when he must have been 75, saying that it was 50 years from his first class. He taught me that what you need to be a teacher is knowledge and passion and commitment. He said, 'It doesn't matter whether you stand on your left foot or your right foot, or it's a small class or a big class. If you have that knowledge and passion, you will teach.' I accepted that, not just at face value but with some thought in it, and it has enabled me to deal with the educational enthusiasms that sweep over the teaching scene from time to time, trumpeting the value of some technology or a particular way of presenting knowledge to the student. These ideologies come and go. What you need to be a teacher is to have a passion and communicate that passion, and the kids do respond. You can stumble and mumble but they'll still respond if the passion is there – and the knowledge, of course. They won't let you hoodwink them.

In those first nine years how did the research evolve?

During the years at New South Wales I shifted from the parallel processing model to study development. I became more and more interested in the ontogenetic development of the brain and the fascination of how the organ could wire itself together. That included a period of sabbatical at Yale with Pasko Rakic, which was very influential: I found him a very powerful, thoughtful figure. But I got stuck into the teaching and balanced the demands as best I could, and we had good, productive years. I must say, looking back, that not quite such a theme developed from them, although the themes have started to develop in the last three or four years.

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Building up research

In the 1980s you got a personal Chair. When did you go on to become Challis Professor of Anatomy?

That appointment was in 1987. It involved moving from the University of New South Wales to the University of Sydney. The department there needed building up, particularly on the research side. My predecessor – Michael Blunt, who was English by origin but had been in Australia for many years – had been an enthusiast for teaching and had introduced two things which I still cherish. One was a commitment to small-group teaching: if you have to have a structural philosophy that's a pretty damn good one to have. The other was to leave a teacher with a group for a long period so they could establish a relationship. We needed building up in research and also in another area. He had done a very good job on classical, topographical anatomy but anthropology, which was the great science in Sydney Medical School at the turn of the century, had languished. An important collection of Aboriginal remains had been built up, and a long string of very important papers on the anthropology of indigenous Australians had been published from that. Another predecessor, Macintosh, was the last of the great figures in that tradition.

So you became curator of these remains?

Yes. And with help, through a generous benefaction from his widow, we refurbished that museum magnificently. We have brought it back to life. Anthropology is the history of anatomy and it's really exciting when you realise that on a bone that is millions of years old you can see, drawn out on that bone, the same tubercles and tuberosities that are on your own, drawn by the same muscles, but millions of years ago. There has to be a sense of understanding of that history, and I want to communicate that to the students. So you can see that I would resist – and I do, but only in the gentlest of ways – the modern philosophy of teaching medical students related to clinical problems only. They will learn about the clinical problem, but they need to experience that scientific and historical background of anatomy too, although we don't really get time to go into Australian anthropology.

But that legacy also brought its burden. Because of the fierce debate – which we had to have – over ownership/control of these remains, I have had to deal, in as deep and analytical and thoughtful way as I can, with the Aboriginal claim to control all bones of Aboriginal origin, and work through it in a sympathetic way. I have as clear understanding as anyone can have, because I come from a victim group. Not in this continent: it is curious that when I travel to Europe I belong to a victim group; when I come back to Australia I am a member of an oppressor group. Sometimes, I tell you, when the shooting's done it is easier to be a member of the victim group.

I have done everything I can to fit in with the Aboriginal claims and always will. But I must say that the Aboriginals and all indigenous peoples claim – absolutely rightly – to be part of the human family and to deserve recognition. But if that is true, then their history is also my history, because I am part of the same family. So the claim of the most extreme to exclude people like me or white anthropologists from the study of these bones seems to me to be flawed. And I am sure that will come to be accepted. I hope that we can with care and responsiveness work through to that position, but it will take a few years.

Being a head of department was a good and bad experience. You realise how much management is required in an institution like a university. I was trying to live up to the old model, in which a professor took some of his or her time out from scholarship to build up the discipline but then got as quickly as possible back to the lab, but I found the universities in a mood of requiring more and more management. They didn't want professors or associate professors to be getting back to the lab; they wanted people who were happy to stay sitting at the committee table and to deal with the flow of reports that the modern institution needs. I stuck with it perhaps a little longer than I should have, but in 1992 the point came to get out. I got out still full of energy and by then it was time to become trained in molecular work. I took that opportunity and have had a wonderful year since. I am looking forward to the next 10 or 20 years.

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Blindness: the ontogenetic link

Brain development sounds fascinating. Was that a rewarding line to take?

Yes. I have realised in the last few years how rewarding, because in that time it has been going better than I deserve. It wasn't a dry patch, because we published paper after paper and I was interested in it, but I had to get the concepts, the data of development into my mind. I had to understand not just the neurones of the brain, to which Peter Bishop had introduced me, but the neuroglial cells, the supportive cells, because they are the architects, the scaffolds of development. And so I spent several years getting into understanding those glial cells and getting the techniques to study them. Introducing one talk, just off the cuff I said, 'Look, techniques give you power but they're also prisons.' I was thinking of this whole mount preparation, which I'd always studied with stains which showed me neurones. I'd worked at it, I'd walked across it this way, I'd walked across it that way. Now we have techniques that show us the glial cells and they break out of the prison of that old technique, but of course only to admit you to another room of the prison. But nevertheless years and years of fruitful work opened up and we began to understand the conversations that glial cells have with themselves and with neurones, and the roles they play.

Then we began to understand the interaction of neuroglial cells with the stress of birth. That is the concept I am now developing, that as the brain comes on line it goes through a period of stress, which is related to its enormous oxygen demand. It goes through episodes of hypoxia, of lack of oxygen. We became tremendously interested in the role of oxygen levels in control of events – of neurone birth, death, movement. Just to give you a feeling for the fruits that have come off the tree: there exist a series of human blindnesses which result from the death of the photoreceptor cells of the eye. Our contribution there – and we are still in the midst of the excitement of it – has been to pick up that the degeneration sets in in this birth period of stress, and realising that if we can relieve that stress we can rescue the cells and prevent blindness.

So there are categories of blindness that can actually be rescued?

I believe so. We have done it in animal models where the gene defects are probably like the human ones, but we have been lucky to get a transgenic mouse in which the condition has been produced with a human gene. If we can save this one – and we haven't done it yet; that's the gleam in our eyes – then we can go through the difficult and demanding process of transferring it to the human models. We can go to the people who handle the human condition and say we've got something to offer. So that adds a medical component to what has been a very basic scientific thrust.

Was it that kind of work, with its genetic aspects, that took you to Jerusalem? Or did Jerusalem actually show you that avenue?

I went to Jerusalem because in 1992, when as I said it was time to break out of the administrative shackles, a group there published the right molecule: evidence of a gene which turns on in hypoxic conditions and makes blood vessels grow to supply oxygen. I wrote to my colleague there, Eli Keshet – he didn't know me from Adam and I wrote by mistake to his graduate student, whose name was first on the paper, but I got the reply from the right guy – and now Eli and I have a close and warm relationship, and I hope to see him in a few weeks to push it along.

So I found myself in Jerusalem. Margaret and I had accepted a period of months apart for me to do this, but I have family there so there was an element of going home and now I have very warm relationships with that university. I had deliberately spent a year there in 1966–67 (Margaret came with me) just to experience that part of the world, and I found myself in the middle of the Six-Day War. I saw Jerusalem under siege and under fire, and even came under fire myself in that medical school.

They were bright and vivid memories but they were already 25 years old. I didn't go back for those memories; I went back for this molecule. We published papers together of which I am very proud. Of course, Eli's input to them was determinative. What pulled me in was the publication of genes which are hypoxia-induced, and I knew that hypoxia was driving developmental events. He gave me an insight into the molecular changes, and that has been of tremendous importance.

You have brought with you some wonderful, beautifully coloured illustrations.

These are a mixture of technology and ideas. The reds and greens are the dyes that work so well in the fluorescent microscope and therefore in the modern confocal microscope, so the technical excitement for me has been to learn to use this superb new instrument. The red is showing the glial cells in a piece of nervous system, the retina. The green is showing blood vessels. What we are exploring in these images, which you see in these red blobs, is the death of the glial cells so close to the vessels. And where that death occurs actually along the vessel, then the vessel breaks out; in this case out of the retina into the vitreous humour of the eye. That is the very beginning of the damaging proliferative vasculopathies which destroy vision in so many people with severe diabetes and venous occlusive disease. With that additional insight I think we then have ways of rescuing those cells, of healing the retina that has been exposed to hypoxia in the diabetic, and better ways than are now available of dealing with these blinding diseases.

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A wider arena: Alzheimer's disease

You have also been drawn into the great and fascinating sink of Alzheimer's disease –being pulled, perhaps, into a wider arena.

Yes. Alzheimer's disease has such links and then such differences. It has been with us for a long time but it has been identified for 70 or 80 years and has become a basket group of dementias for which we don't understand the cause. Alzheimer's thrust, when you read his original papers, was to distinguish his patient's dementia from a dementia that arose from syphilis, which we don't see any more. He drew attention to two pathologies, the plaques and tangles prominent in the cerebral cortex of the sufferers. Molecular biologists today have done beautiful work on both of those molecules, and on another protein, apolipoprotein E. But I was drawn in first through the Hebrew University connection and through a Jewish businessman, John Hammond, in Sydney, who was very committed to this because of his wife's affliction and who felt his own strength failing. He is in his late 80s now. I had returned from places that made sense to him and he asked would I be a trustee of the Sir Zelman Cowen Universities Fund, a charitable trust which he had established. That drew me into the field, and to understanding it.

A paper in a major meeting in Kyoto last year argued that the best correlates of Alzheimer's disease are not plaques and tangles but brain inflammation. I knew enough of the glial cells of the brain and the cells which might be involved that I could read the data – and it had an element of the loner idea which maybe struck a chord with me. I came back from Kyoto with this idea that I felt had fallen on a mind that was ready for it (but I don't mean that immodestly) and convinced that it was worth exploring. Not that it was true, because science isn't like that, but obviously the other techniques were not offering anything to the patient, the sufferer, and this did. There was evidence from retrospective epidemiological studies that anti-inflammatory drugs could prevent Alzheimer's disease. There was one prospective study which suggested that once it had started you could stabilise it and stop it from progressing. And those drugs are available. They all have side-effects, but it's so much more than is available to the sufferer.

So I persuaded the trustees; we convened a group of scientists to monitor it; and we convened a working party of real Alzheimer's scientists who took it on board. We have funded it and it is about to be launched officially. We aim to report within 12 to 18 months, and I hope we will do enough work to convince our Medical Research Council that there is a case for clinical trials. If so, it's exactly what that Fund wanted: at the earliest possible moment to produce something that could offer some hope.

It must be nice for you to have made the transition from focusing on nerve cells and the structure of the brain to dealing with wider human need.

Yes. But certainly it has a tie-in with the work I am doing now on cell death and survival in development, because we are talking about massive neuronal death. I believe it's quite different. I don't think it is driven, as it is in retinal development, by hypoxia or oxygen radicals. It may be an activated microglial cell rampage that sweeps through the temporal lobes and then the parietal lobes, and destroys the brain of these sufferers. And I think if that is true we can stop it. That would be a tremendous gratification.

We are beginning. We have funded four projects, one of which includes a preliminary clinical trial. That will be carefully done because the drugs have side-effects. One of the valuable contributions of the molecular work is that it has identified a group of people who are really likely to get Alzheimer's disease and to whom you have some justification to say that they really should take on this drug, with all its side-effects, if they get to a certain age. There is a great deal of caution about clinical trials. I am not a clinician but I am of the view that you have to try, otherwise you are never going to break down that caution. I am hoping I will see strong methodological connections that might bring me to work on Alzheimer's but I'm content at the moment to have coordinated that group and I'm looking forward very much to helping write the report.

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And wider still: animal welfare

Jonathan, we've taken you through the central strand of your career – neurophysiologist, neuroanatomist – to some new things at the cutting edge. This wider holistic person has published in a number of other fields as well – concerned about animal welfare, animals in experiments; concerned about population and the future of the world environment, and a number of other issues of public debate.

There were two or three of those debates into which, yes, I was drawn – by circumstance and by inclination too. I was drawn into the one over animals by a rising debate, principally in the United Kingdom but with its eddies coming out here, over vivisection. It was a renewal of an old debate and I wanted to get to the bottom of it. Having done a lot of animal-based research, I wanted to understand the moral challenge and respond to it by changing or not changing what I did. I came to understand the nature of debate, the tricks and also the earnestness of debate. I came to learn a lot about how well-meaning, good-hearted – and I mean that without patronising – societies like the British Union for the Abolition of Vivisection could be raided by fascist societies, as they have been, how anti-vivisectionism could become a cloak for a general anarchism, and how to deal with that, how then to respond in the most creative way. And that was obviously by trying to clear away the terrible edges of that debate and concentrate on the core moral issues with the real and earnest people.

That had its awful moments. The Australian group drew from England in deciding to focus on individuals and for some reason I turned out to be their focus. I decided not to delist my telephone but to just walk out and take the flak. It drew me into public debate and also into earnest writing, and I've tried to do both of those as best I can. I think we have managed, with a lot of input from a lot of people, to take that debate in Australia along very creative lines, where the issues of animal experimentation are handled by institutional ethics committees in which the debate takes place on a weekly or monthly basis, in detail, with animal welfare people brought in. And now there is legislation through all the States. So yes, I played a small role in that: some of my writings went beyond what was in the literature and maybe I made a contribution.

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Population biology

How were you drawn into the population debate?

That was really through my Academy work. In the '80s I was privileged to be asked to be Biological Sciences Secretary for the Australian Academy of Science and had a thoroughly enjoyable and instructive four years seeing science, as academies do, as groups with energy needing to be brought together. I was very encouraged by that fascinating period.

I was asked to represent the Academy at an international meeting of academies which was held in Bologna in '89 with the theme 'Scientific Problems of the Next Century'. Sitting down with a blank page to prepare a presentation, it slowly became clear to me that overpopulation was the great challenge we faced. It has a lot to do with medical history because the reason we are overpopulated is the success of medicine in reducing death rates. When I got into that debate I found it quite fascinating, because I began to disagree with the David Suzukis and the Paul Ehrlichs of this debate – not because they weren't right on so many of the issues but because they constantly took the high moral ground. I felt they didn't deserve the high ground and that taking the high moral ground was destructive.

Despite both Suzuki and Ehrlich being biologists, they have not dealt with the reality that we are overpopulated not out of carelessness or mismanagement, as is their constant refrain, but out of our success in doing something which we all accept is good, and that is keeping children alive. That although birth rates have fallen since the middle of the 1850s, the fall in death rates has been so dramatic that we have massive overpopulation. And if that is true, then the evil of overpopulation that is making a mess in so many ways is deeply entwined with the good of what we do. Until their entwining is realised, they can't be untangled.

I like the start of your lecture. You went back to Dickens: 'It was the best of times, it was the worst of times.'

Yes. It is the human condition that the good and evil of what we do are tightly bound together, and those who would preach usually are thinking of themselves as having the insight or faith to separate the good from the evil – which they have not. You can sense in that a loner's position, that I was not a conservationist nor a populationist nor a growth advocate, but I felt that I had identified the core issue of overpopulation and that it was worth arguing. People do respond to it.

The next step in that path, after the talk in Bologna and the Australian Foundation for Science lecture in 1991 in Adelaide, was to convene a symposium – which again the Academy was generous in taking under its auspices, not as its opinion but by giving it its platform. We were very lucky in that we met in 1994, just as the Australian Government through Barry Jones' committee (the House of Representatives Standing Committee on Long-Term Problems) was convening its inquiry into the population of Australia, and our submission was just in time for that. I think we made an important contribution there. That was a rewarding experience.

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Optimist or pessimist?

You're a great private crusader, but these are tremendously big issues and I don't see them all being untangled in time. Are you an optimist or a pessimist?

It's difficult sometimes to be an optimist. But in that lecture on population control I drew an analogy with the Cold War. Underlying what was happening during the depths of the Cold War – mutually assured destruction (MAD for short), the constant building up of the arsenals – was a constant series of meetings and a series of treaties on the use of the seabed, on nuclear weapons in space; the START I negotiations go back, I think, to the middle 1970s. START II is a bit later. Good was emerging but it was being outpaced by the growth of the armaments. And so I thought it was a striking and unique moment in history when Gorbachev declared that the Soviet Union would not again intervene militarily in Eastern Europe. It was an absolutely key moment, when a very proud revolution announced its limits. I can't remember another revolution that has done that quite so superbly as the Russian Revolution, and the Russians deserve enormous credit for saying, 'This vision has come to its limits.' That opened up so much, leading to the first disarmament since the Romans formed the centurion brigades. It's been an upswing ever since.

So yes, in times when we can't see the solution to a problem such as overpopulation we must work to lay the framework, because somewhere it will come. There is some element of optimism, in that birth rates are falling so universally that the United Nations people are correct to project a levelling of the population. It is at a very high level, but perhaps somehow we can manage through to there and learn not constantly to grow the population. I think we are going to accept the limits to population growth very soon, even within 20 years.

Limits to growth are going to be harder to accept. Governments who accept that their population should stop growing – I think the Australian government is close to doing that – will not be ready to accept that economic growth should stop, because it is the solution to unemployment, to the problems of the needy, of how to bring the needy help without soaking the rich. That second stage, of stopping economic growth, will be much slower. I can see reasons for optimism, but no reason for stopping developing the infrastructures with which we must one day solve these things, in good times or bad.

Your exciting book Parallel Processing in the Visual System, published in 1983, seems to me to exemplify your work. Such a contribution to the literature was overdue.

It was due in its time. It picked up on a theme that for me finished in the early 1980s, of the parallel processing in the visual system and its implications. It has in it two or three chapters which distil what I had come to understand of the heuristic debate that underlay both the creative ferment and the conflicts of that time, and have been reference points for me ever since. I tried to make it very strong empirically, because you become vulnerable as a scientist if you spend too much time with the cheerlessness of the philosophy. You've got to be careful to understand the philosophy but then not to be too entranced by it, but I think that scientists do need to understand the philosophical framework, the philosophical problems within which they are working. As I said before, Eccles and Popper were very influential with me. Popper, in one of his books, wrote that if you want to understand a philosopher you should understand the problems he was dealing with. That has helped me a lot, including when I go into the public debates: why are people taking particular positions, what are they defending, what are they trying to bring forward – a very good reference point that has helped me in these many ventures.

Jonathan, we've covered a rich story, a rich tapestry. It's been fascinating to talk with you. Is there anything behind the story that I have missed?

I've been lucky to be born at a time when I didn't have to die in somebody's army; I've had good health and a clear run. I have debts all over the place: to family, to my present family, to scientific mentors. I have tried to name the important ones but I'm sure I've missed some. I am still thoroughly enjoying it. Perhaps I could end on the message that the excitement is there and I'm looking forward to the next 25 years with the same sort of zest.

Before that's up, though, perhaps I'll come back and talk to you some more.

Sure. I look forward to that. And thanks for having me.

For so many things, thank you very much.

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Introduction 

Dr Leanne Armand received a PhD in geology from the Australian National University. Her thesis work focused on the use of algae remains as an indicator of sea surface temperature changes and sea ice estimation. The algae remains were found in sediment cores taken from the southeast Indian Ocean. As an Australian Research Council postdoctoral fellow with IASOS (Institute of Antarctic and Southern Ocean Studies at the University of Tasmania) she developed this work further, especially in the area of estimating sea ice extent in the Holocene and over the last 190,000 years.

Now working in the Biogeochemical Cycles Program at the Antarctic CRC, she investigates biogeochemical cycles using algae collected in sediment traps at certain ocean sites between Australia and Antarctica and also continues her sea ice research. Her work is important in gaining an understanding of how sea ice and sea-surface temperatures vary naturally over time, and how this natural variation influences climate.

Interviewed by Ms Marian Heard in 2001.

© Australian Academy of Science

Rutherford Robertson was born in Melbourne in 1913. He attended Carey Grammar School there and when his parents moved to New Zealand he attended St Andrew's College in Christchurch. In 1934 he received a BSc Hons from the University of Sydney. In 1936 Robertson went to England to study for his PhD at Cambridge University where he worked on the relationship between cellular respiration and salt uptake by cells. He returned to the University of Sydney as an assistant lecturer in 1938 where he worked on apple and wheat storage.

Between 1946 and 1958 Robertson worked at CSIRO as the head of the section working on plant physiology and fruit storage. He spent 1959 as a visiting professor at the University of California at Los Angeles and on his return to Australia joined the Executive of CSIRO. He received a DSc from the University of Sydney in 1961, and in 1962 was appointed professor of botany at Adelaide University, a position he held for 7 years. Robertson moved to Canberra in 1969 to become master of University House at the Australian National University. He then became director of the School of Biological Sciences at the Australian National University, a position he held from 1973 to 1978. Sir Rutherford passed away on 5 March 2001.

President of the Australian Academy of Science 1970-74.
Interviewed by Dr Max Blythe in 1993.

Contents

• Family background and early life
• First interest in science
• Education
• Entering scientific research
• A marriage of scientists
• Activities at Cambridge University
• Teaching science
• Solving food storage problems
• Investigating active transport and respiration
• Overseas studies
• Promoting scientific research

Family background and early life

Perhaps you could start by telling me about your early life.

I was born in Melbourne in 1913. My parents were living there because my father was a Baptist minister and he had churches in Melbourne. My parents were strictly religious people, as you might expect in the Baptist denomination. While my father was busy converting people, my mother, who was well-educated and well-read, had a broader perspective. Certainly I owe a good deal of my early upbringing to her wisdom.

What kind of person was she?

She was very investigative and really got me interested in science, particularly when we settled down after World War One. During the war, my father was a chaplain to the Australian Forces in France and was away for two years. My mother, who came from Queensland originally, took me to Brisbane and we stayed with my grandmother. My father came back in 1919.

By that time I'd succumbed to an epidemic of polio (or infantile paralysis, as it was called in those days). When we went by train from Brisbane to Sydney, I remember limping along the platform to meet my father – a forbidding figure in full uniform – Captain Chaplain Robertson. So those are my earliest memories. We then went on to Melbourne, where he was going to a church in the suburb of Canterbury.

Talking about your family...you have an interesting Scottish and English background.

Yes. The Robertson name comes from a family in the Shetland Islands. The Shetland Islands were in a pretty bad way in the middle of the 19th century, and there was a great deal of emigration. Our branch of the family migrated then and went to settle in Victoria. My grandfather Robertson married another Scot, Miss Cairns, whose mother's maiden name was Rutherford; that's how I got the Rutherford in my name, because it was a family name given as a Christian name.

On my father's side there must have been genes for preachers, because almost every man was a preacher. In the 19th century they conducted the sort of tent-missions which travelled around Australia singing and preaching. On my mother's side my grandfather (whom I never knew, because he died before my birth) was a very successful businessman in Brisbane. That family too was well-read and well-educated, but none of them was encouraged to take tertiary education in those days; they went into business.

Just thinking of that line of preachers – tell me more about your father.

He was a great influence as far as I'm concerned. While he did believe it was necessary to save souls, he had more than just a conversion instinct. In addition to doing theological courses, he studied history and social studies. He preached what he himself would have termed a 'social gospel', which meant that Christian people had to do something for the community as a whole and not just look after their own heavenly futures. That was unusual in those days.

You mentioned being ill as a boy and winding up with a limp. That must have been hard to cope with.

It wasn't easy, though my parents encouraged me to do everything to strengthen my weak leg and to take part in sport as far as I was able. If you recover from poliomyelitis, you seldom have pain subsequently, just weakness, and I didn't find that too much of a burden. The worst part was being dragged around to various people who thought they might be able to cure it. There was always the possibility that the next person I was taken to might have a better form of massage or something of the kind.

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First interest in science

What stimulated your early interest in science?

My mother. She did not have any specialised knowledge, but from reading and from occasional lectures, she became interested in what was happening in science and what scientists were up to. This was in the early 1920s in Melbourne.

And she'd talk to me about it. For instance, I remember when Ernest Rutherford visited Melbourne, she went to listen to his lecture and to learn something about what such people were up to, without knowing any of the details. She encouraged me to have a similar interest in science. When I began to see that chemistry was an interest, my parents didn't buy me a toy chemistry set. Instead, they went to a friend who was an industrial chemist and said, 'What sort of things could a boy do a few experiments with?'

So you built a bench and you got to work.

Yes, that's right. I didn't discover anything very vital, but I enjoyed it.

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Education

And your early schooling in Melbourne?

It was mostly enjoyable. Because of my inability to get about very well, the first school I went to was what would have been called in Britain a Dame's school. It was a school mainly for girls up to about 14, but which took a few small boys. It was very near where we lived, and was run by three sisters, the Misses Hester. I learned not only what I was supposed to in my own class, but because there was really one big room with all ages in it, I also learned what was going on in the other classes. I really enjoyed that period.

Then I went to what was the beginning of a primary and secondary school, called Carey Grammar School, which had a Baptist foundation. I was there for three years before my family moved to New Zealand. I enjoyed my time at Carey, although I began to learn what it was to have hard knocks with people with whom I didn't see eye-to-eye.

Did Carey contribute towards the scientific career that was to follow?

Only a small amount. Science was a subject that was taught, I think, only in the last year that I was there, by which time I was about 12. As far as it went, that added to my interest, but it wasn't very exciting science. Science classes in those days tended to start with mensuration, so we spent our time looking at rulers and protractors and one thing and another. This didn't seem as interesting as chemistry, which I wanted to get involved in.

In the mid-1920s your family went to New Zealand for your father's posting?

Yes, it was a fascinating experience. We went to the South Island of New Zealand, to Christchurch. The advantage of the South Island from my point of view and from my parents' point of view was the proximity to the Southern Alps. I had several opportunities to go to Franz Josef Glacier and to the high mountains in the vicinity of Arthur's Pass, which was one of the ways through the mountains in those days. That was a good experience.

What kind of school did you go to?

It was a Presbyterian school, a boarding school for sons of farmers, but I went as a day boy. The social life was fairly demanding and enjoyable. I had to turn up on Saturday afternoons to watch the first 15 play the rugger match, and I had to take part in sport, which I enjoyed to the best of my ability. And I think the teaching on the whole was good. Certainly the ethical outlook of the school was very good, and I enjoyed the teachers I was associated with.

Do you remember anyone in particular teaching at that school?

Well, I remember the mathematics teacher, Leadbetter, who was very good indeed. He was something of a hero to the boys, because he was the sprint champion of New Zealand at the time he was our teacher. And in those days, anybody who could run 100 yards in 10 1/5 seconds was automatically a hero.

Leadbetter went on to become headmaster of one of the big and famous New Zealand high schools. I think he went to Waikato, and he then did an unusual thing: on reaching retiring age, he took holy orders in the Presbyterian Church and became a minister. I was pleased to call on him in later life when he was a retired preacher.

And what about science teaching in the school?

The science subjects were mostly restricted to chemistry and some very elementary physics. We didn't really have a scientist as a science teacher. I remember that I had a little trouble when it was said (and I suppose I'm talking about 1927 or so) that we were now going to talk about the atom, which was the smallest indivisible particle – that there was nothing smaller than an atom. And I said, 'But, Sir, hasn't...?', and went on to suggest that someone had split the atom. The master at the time was embarrassed and said, 'Yes Robertson, come and see me afterwards'. So afterwards he said, 'You're quite right, Robertson, but that's not in the syllabus'. So that was about the level of science we were getting at that stage. But I didn't lose my enthusiasm for having more.

You came to Sydney next, and went to university there?

Yes.

It must have been quite difficult going to Sydney University from that kind of background.

Yes, it was, and in some ways it was a mistake. My father, consulting me, toyed with the idea of my going to a school in Sydney for another year or even two. I was only 16, but I had a matriculation qualification, which Sydney University accepted. A friend of the family who was a school teacher said, 'Why don't you let the boy go to university? Even if he fails and repeats, it won't matter very much, and it would be better for him to start getting university experience.' In fact, that wasn't quite so, because I was ill-prepared and had to work so hard to keep up. I didn't get much chance to grow up – I was too busy working in the first year or two. That all changed with a couple of years' maturity.

And what did you study?

Chemistry, botany, physics and geology in the first year. Then it split down to more chemistry and botany, and I did zoology as a first year subject in second year.

But by the time you got to third year, you were really beginning to find that botany was coming out on top?

Yes, that's right. The botany that I was doing then – under the leadership of Professor Osborn who subsequently went from Sydney to be Professor at Oxford – was more interesting than chemistry. It was the way it was taught; that was largely because there was a lot of memory work in chemistry, which I found less challenging than thinking out the solutions to problems that I was getting in the biological courses. Subsequently I made up for that, because my interest in chemistry didn't decline. I learnt much more in graduate years than in the undergraduate years.

And Osborn?

He was a very good lecturer, sparkling and lively. His lectures were appreciated by first year students. He was not so good for third year students, because he didn't give us as much of a feeling for the research atmosphere then as successful teachers of senior students do now.

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Entering scientific research

In your fourth year, you had an option to do a research year?

Yes, that's right. I chose to do the honours year in botany, which consisted of a great deal of reading – a long reading list – over the whole range of the subject, including its history. Osborn suggested the research problem, and then left the honours students alone to work out what to do, though he gave advice from time to time.

But he set you a rotten problem?

Well, it was partly my own choosing. It was to work on the way the pores in leaves, called stomata, open and shut. It was quite fascinating, and I did manage to get a few results. In fact, the fourth year students in the different subjects were invited to compete for a prize based on the thesis or paper they had written, and I got a student prize for the work that I was doing. I also got a first class honours degree, which showed that I was enjoying it and making something of it.

Looking at that work, you made all kinds of apparatus.

Yes, one of the physical chemists helped me. I learned how to make apparatus by blowing glass. I did that primarily so that I could analyse for carbon dioxide and oxygen, which we believed going in and out of the stomata had something to do with controlling their opening and closing.

You had to make a high vacuum?

Yes, in attempting to get analysis for very small quantities of gas. I didn't use it throughout the research, though, because I found I could get rather bigger volumes of gas than I had been thinking of. I then made an apparatus which was based on the Haldane apparatus, which was a gas apparatus. That worked for what we were doing at the time.

But you didn't resolve the problem you initially set out to resolve?

No, so I talked to Petrie in Adelaide about it during the summer holiday. I should explain that this was a plant physiological problem and there were no plant physiologists in Sydney at that time – none at all. The only two of any stature were Petrie at the Waite Institute, University of Adelaide, and Wood, who was Professor of Botany, also in Adelaide. I got an opportunity to discuss the problem with Petrie, and he made some suggestions, which I worked on. But they weren't terribly successful either, at least not in my hands, with the equipment available at that time. But the important thing about the contact with Petrie was that he said, 'If you want to be a plant physiologist, you must get to Cambridge.' This was good advice.

How did you meet Petrie?

My father took a busman's holiday by going and preaching every Sunday for a month in a church in Adelaide, so we were living in Adelaide. He did that in two successive years. On the first occasion, through friends of his, I was taken round the Waite Institute and met Petrie. And Petrie, being a Sydney graduate, took an interest in me, and we had some discussions. The next year my father was going over again, and Petrie said, 'Why don't you come and work in my laboratory for the three or four weeks that you're here.' That was a great experience in itself.

How did you find Petrie?

He had a very sharp intellect and was very critical – so critical at times that he offended the people that he was giving helpful, critical advice to. But I got on very well with him.

From there you went to Cambridge?

Yes, I got a scholarship. Some of the money that was raised in the Prince Albert Exhibition of 1851 was used to set up scholarships called the Exhibitions of 1851. These scholarships were mainly responsible for taking colonial or dominion students to England. Most of the students went to Oxford, Cambridge or London University, but not necessarily. When I was fortunate enough to get one of these, I went to Cambridge on Petrie's advice.

I was wondering whether that's where the term exhibitioner had its origins.

I think it might be. The list of people who went and became famous since is long. I think Rutherford had one of those scholarships when he first went, and certainly Mark Oliphant had one, going from Adelaide in his case.

So you went from Sydney to Downing Street and St John's College. What a botany school! When did you arrive there?

I arrived at the beginning of October 1936. I went in to see Briggs, of course, and gave him the work that I had been doing on stomates. He was not too uncomplimentary, but it was obvious that he had something that he really wanted his next research student to do. He suggested that I might put this stomate work aside and work on what was really interesting to him. That seemed to be good policy – to do what your supervisor wanted done – so I had no difficulty in doing that and was completely fascinated with the problem.

What work was that?

The best way to start is to say that all cells – plant and animal – take soluble constituents (nutrients and so forth) into them. They do it in such a way that quite often these constituents are being moved against their natural tendency, against the concentration gradient which makes them diffuse back out, but there is something that spends energy on taking these constituents into cells. Not at the time I started with Briggs, but later on, this came to be called active transport, meaning active in the sense of work having to be done to make these things move. We called it accumulation in those days, because it resulted in salts such as sodium chloride or potassium chloride accumulating in plant cells. If some work is being done in living organisms, the energy for that work comes from the process of respiration.

Briggs invited me to combine the measurement of the respiration with the measurement of the salt accumulation. We used slices of ordinary carrot roots, which were living cells. As the salt went into the tissue, the electrical conductivity of the solution that it had gone from would decrease because there was less salt in it. We had electrodes in the external solution, and we could measure the amount of salt that was going into the carrot discs.

The respiration resulted in carbon dioxide being given off, and we developed a technique for measuring the amount of carbon dioxide given off, which also depended on electrical conductivity of a solution. We bubbled air past the respiring tissue, and the carbon dioxide given off was taken into the air stream. We passed it through sodium hydroxide, in which the carbon dioxide combined to make sodium bicarbonate. The sodium bicarbonate has a lower conductivity than the sodium hydroxide, so we were able to get conductivity shift again. So it was a double conductivity shift that we were looking at.

Did it correlate the respiratory rate with salt uptake and accumulation?

Yes, it did. The system we were working with was very advantageous, because after we cut the tissue and put it in aerated water for 24 to 48 hours, the respiration rate dropped very low. When we put salt on it, it rose again and – to make a long story short – the increase in respiration was proportional to the amount of salt that was taken in. A good deal of my work went into establishing that close relationship with aerobic respiration.

And that was the core of your thesis?

Yes. I finished the thesis in December 1938, and I was examined orally by Maskell, who was a famous physiologist and my internal examiner in Cambridge; and Bennet-Clark, another physiologist who was the external examiner. They give me a viva, as it's called. Nobody told me whether I'd passed or not – nobody at all. There wasn't a hint. I was on the ship going to Australia, somewhere near Cape Town, when I got a telegram from my tutor in Cambridge saying that my PhD had been passed.

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A marriage of scientists

I want to go back a bit before the doctorate to your marriage to Mary Rogerson. Could you tell me about her background.

Yes. My wife is of Scottish parentage. Her father came out to settle in Australia in 1911 I think, and his fiancée came a year later. He sold a farm in Dumfrieshire, which had been in the family for a long time. The name Rogerson is quite famous in that area. They were either farmers or doctors. Indeed, there was one John Rogerson who was medical adviser to Catherine the Great because of the connections between Scotland and the Russian court in those days.

When Mary's father came to Australia, he bought land first near Canberra at Gundaroo, and then in 1920 at the place where we now live – Binalong – which is about 95 km from Canberra. Mary was a science student at Sydney University when we met, and we were engaged to be married before I went to Cambridge. We were then facing a separation of a possible two years, but fortunately, with some financial help from our families and finding that I could do better with my scholarship than we feared, she was able to come over at the beginning of the '37-'38 academic year. We were married in September 1937 in All Saints Church, Jesus Lane, Cambridge – which sounds great.

And your marriage has lasted all these years?

Fifty-five years and still going strong. Mary has been an enormous support. In our Sydney days she used to do part-time demonstrating, which was practicable. We had a son who had to be looked after, and a full-time job would have been difficult. She continued that interest until we went to Adelaide, when we made a choice of our own that perhaps a professor's wife being on the same staff might not be a good idea. That might have been an error, looking back. She could probably have continued demonstrating part-time there, too. But she not only helped me with experimental work from time to time, but also edited a great deal of my writing. She criticised my papers, and did all the typing before word-processors came in.

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Activities at Cambridge University

I'd like to spend a bit more time talking about Cambridge in those years.

Cambridge was a very exciting place in the 1930s. The Cavendish Laboratory, with physics led by Rutherford and a collection of stars around him – Cockroft and others – was doing exciting things that were reported almost every month. It was a great privilege to be able to go and listen to the great Rutherford give an occasional lecture. Plant physiology, of course, connected up with biochemistry, and the biochemistry school under Sir Frederick Gowland Hopkins was especially strong in those days. Among the people that I knew there were Dr Joseph Needham and Dorothy Needham, and Robin Hill, Pirie (who worked on viruses), and Dick Synge (who subsequently won the Nobel Prize for work on paper chromatography in collaboration with others).

And Briggs?

Briggs was one of the most imaginative plant physiologists that I met. He placed a strong emphasis on the importance of physical chemistry and mathematical analysis to the basis of living processes. He was highly critical and appeared offensive when not meaning to be. It's said that once, just before my time in Cambridge, he was asked to comment on a lecture given by the Professor of Botany from either Edinburgh or Glasgow – I'm not sure which. The professor was a very distinguished figure in his day, but in his older years, he wasn't presenting things with the rigour that they deserved. Briggs said so in the audience. The gentleman, when asked to reply to Mr Briggs (as he was then), said that he didn't think he wanted to waste his time on some up-start of a young man who didn't know what people were doing. The story goes that the poor chap wouldn't eat his breakfast the next morning. He was staying with Sir Albert Seward, and Lady Seward had to ring Briggs and suggested that an apology would be appropriate. Briggs was very upset about having done this; he had no idea that his trenchant criticisms were in any way unfair, and he had no idea that he was upsetting anyone. When he made those sort of criticisms, they would be pretty sound.

Briggs was very supportive of your work...

Yes, he was one of the two great influences in my career.

And so, Cambridge. Cambridge was under the influence of the Spanish Civil War at that time.

Yes, it was. In 1936-37 we were concerned about the way things were developing in Spain, exemplifying, a lot of people thought, what was going to happen in Europe generally if Hitler cut loose as he was threatening to do. And, as a person interested in the social relations of science (perhaps due to my father's influence), it was inevitable that I took an interest in the sorts of groups that had that in mind.

One of these was the Association of Scientific Workers, which tried to get scientists to acknowledge their social responsibility and the contribution that science could make. I became secretary of the Cambridge branch. At that time, not surprisingly, such bodies tended to attract a number of people who were on the left side of politics, including some people who were communists – a few card-carrying communists, others secret communists – and I found that atmosphere very interesting. I was never a communist myself, because I tended to be suspicious of things that some of these people took as acts of faith.

The other body that I was associated with was the Cambridge Scientists Anti-War Group, and this was the one that related particularly to the way things were developing in Spain. We were concerned that the conservative British Government was using some technical knowledge to prepare people for the possibility of Britain being invaded – like advice on how to gas-proof rooms in houses, should there be a gas attack as part of the war. At that time we published a little book called The protection of the public from aerial attack.

You were a contributor?

Yes, I was one of the junior contributors. Somewhere down here in the list of the editorial committee, it says R.N. Robertson BSc. This gave me an interest in meeting a lot of people that were seriously concerned about what was happening. At this time the great J.B.S. Haldane, an acknowledged communist, was writing daily articles for the communist paper, the Daily Worker. He gave an unfavourable review of the book, which surprised my colleagues, because they thought that Haldane should be on our side. So I was deputed to go and see Haldane, who was in London at that time. I called on this great man, who really was a great scientist. He was just back from helping the government in Spain against Franco, and he gave me a very good hearing. He agreed in the end that in one of his criticisms he may have been right, but in the other he was wrong and he apologised. So, as far as I was concerned, that was a good experience.

I have to say that I was surprised at the devotion of some of the people on the left, including Haldane, to things that obviously were not as rosy in the USSR as these people believed. As far as I was concerned, when Hitler and Stalin made a pact, that was the last straw. I couldn't believe this stuff anymore, and was surprised that some of my friends remained with it until later years. Most of them were put off by the invasion of Czechoslovakia, I think.

So that was the atmosphere of Cambridge, and one of my personal experiences relating to the atmosphere at the time. When summer came in 1938, I went to Leiden for a couple of weeks for further experience in glass-blowing.

That was something that you really got into?

Yes. Knowing that I'd have to make equipment for myself when I got back to Australia, I was keen to do that. After Leiden, Mary and I went to Germany, but with some misgivings, because by this time it was early September 1938. We went to Munich, because a friend of ours was working there, and we were surrounded by Nazi uniforms. We had a very pleasant visit, except when we got into an argument with one of the German soldiers in the Hofbrauhaus. We had a friend with us who was fluent in German, and it was quite clear that what they thought they were going to do was very different from what we thought they should do.

We returned to England in that September, and under strict instructions, the first thing we did was go to the local school in Cambridge and collect our gas masks, because it was thought that war would break out within a month. As it happened, Chamberlain went and talked to Hitler – what became known as the Munich crisis – and the war was put off for a year.

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Teaching science

And so you returned from Cambridge to Sydney University and the Botany School you had left...

That's right. Osborn had left and early in 1938 Eric Ashby, who subsequently became Lord Ashby, had taken over the professorship. This revolutionised the outlook and the way things were done. I went back to Sydney University as an assistant lecturer. Ashby gave me responsibility for the practical classes in plant physiology and some of the first year practical classes in botany, and this was a good experience. First of all, I found that I really enjoyed teaching. Secondly, Ashby said that the important thing about teaching is not to give students a lot of facts but to make them think, which suited my approach to science. Ashby encouraged us to make every practical class like a mini-research project, where the students were encouraged to investigate for themselves, draw their own conclusions, and discuss it with us when or if they got stuck. I enjoyed that...

But different to your time as a student.

Yes, it was. It was different from organic chemistry, which was all memory work, and from lots of botany, which was also memory work.

So you settled in well?

Yes, but then the war came in September 1939, after I'd been back for about eight months. Ashby was called on to do war work. Towards the end of the war he was asked by Australia, with the connivance of Great Britain, to go as a counsellor to the Australian Embassy in Moscow to further relations between the allies and Russia. By that time the USSR was fighting Hitler.

I was put immediately on the reserved occupation list, which meant that I couldn't leave. I took over some of the things that Ashby had been doing, even while he was still in Australia, because he was busy setting up a Science Liaison Bureau.

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Solving food storage problems

I also got involved with CSIRO, with problems that arose out of the war in relation to two food products. One was apples. All of a sudden there were no ships to take our export apples from Australia to other parts of the world. We needed to store them for as long as possible to spread the marketing in Australia, so that there'd be minimum loss for the growers. We worked on things like oily layers on the surface of the fruit to try and restrict the respiration and hence make the fruit live longer.

The other food product was wheat. There was so much wheat that every silo in Australia was filled, and there was nothing to do except put water-proofing on the ground, make a huge stack of wheat and then build a roof over the top. They still do it these days with excess wheat. The stack would be about 10 feet deep on the edge and 40 feet deep in the centre, and the size of the structure would be as large as 5 or 6 tennis courts.

The problem was that the wheat was getting hot. It was racing up to 40 degrees centigrade, and the question was, 'Why is the wheat getting hot?' Dry grain has a very, very low respiration rate, and we didn't think it could produce enough heat to do this. With the CSIRO people, we discovered that there were grain borers and grain weevils boring into the wheat stack, and it was their respiration that was bringing the temperature up. So, those were the two things that I was doing part-time.

You made quite a contribution to apple storage, though, didn't you?

Yes. That led me on to what happened at the end of the war, which was a change of jobs. In 1946 I moved over to head a section of CSIRO. That gave me staff who did experiments on the practical side, like how to store fruit. It also gave me some extra research assistance for continuing the work that I was doing in fundamental ion movement and respiration, which was encouraged. That was a very happy period, especially as CSIRO agreed to my continuing to run the practical classes in plant physiology at Sydney University. So I had the best of both possible worlds. I did that until 1958.

And you confirmed something about fruit storage that was important?

Yes. With the staff that I had in the fruit storage work, there were experiments going on in cold-rooms, cold-rooms with modified atmospheres and so on. I had very competent people to do this routine work. But, in the physiology of fruit, we followed up an observation made by Martin, a CSIRO colleague in Tasmania, that large fruit could be of two kinds: one kind would keep reasonably well in storage, but the other would keep badly.

We were able to show that if the fruit grew large because it had produced a lot of cells (in other words, there'd been a lot of cell division) but the average cell size wasn't very large, that fruit was a good keeper. If the fruit grew large but had produced few cells, so that the average cell size was large, that was a bad keeper. This was important information. We sent advice to growers and people storing fruit about how to divide their batches of fruit to keep the bad storers for a shorter time.

This would have had widespread economic importance.

Yes. Martin had most of this information, but he asked us to find out the explanation for it. So that was the fruit storage work. At the same time we were doing work to try and explain why or how the respiration rate of fruit changes and the time it needs for ripening – technically called the climacteric rise in respiration. We tried to determine what controlled the climacteric rise, whether it might be due to the balance of the free phosphate and bound phosphate in different constituents of the fruit. If there was a lot of free phosphate, there was plenty of opportunity for the respiration to rise. If the phosphate was bound, the respiration was going as fast as it could, because it couldn't send compounds away.

Was that a good hypothesis?

Yes, except that in the end, like a lot of science, it didn't turn out to be universally true. There were some varieties of fruit for which the explanation was different.

Was it true for apples?

Yes, it was true for apples, but it became more complicated to fix all the details.

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Investigating active transport and respiration

And at this time, you were still excited about the active transport?

Yes, in the 1950s we were investigating the active transport and its relation to respiration at a more biochemical level. Respiration produces energy, but it distributes the energy through a phosphate carrier that is called ATP (its longer name doesn't matter for our purposes). We found that if respiration was proceeding and the active transport was proportional to the respiration, the respiration could be knocked out with various inhibitors and the active transport would be knocked to zero.

One of the things we found was that an inhibitor of the formation of ATP would also knock the active transport out. That was a very important observation, because it led on to later aspects of my work that were to connect up with the brilliant work of a man called Mitchell. But the way in which the respiration was affecting the ATP made us wonder whether the active transport was directly dependent on the respiration, because we could knock it out with respiration inhibitors; or whether the ATP needed to be formed first and the active transport depended on the ATP, because we could knock it out with ATP inhibitors. We did a lot of work to separate those two possibilities.

We were bothered about this through the 1950s. In 1959 I went to the University of California at Los Angeles as a Visiting Professor for the academic year, and while I was there, I attempted to follow up the literature in my own previous work, about whether the active transport was dependent on the respiration or the ATP. As early as 1940 we had shown that the respiration and the active transport were stoichiometrically related (that is, the number of oxygens being taken up was quantitatively related to the amount of salt that was being accumulated). At that stage, we decided that it was mainly a respiration feature, and we believed that the respiration was producing a separation of positive and negative charge – the negative charge being an electron going one way, and the positive charge being a proton going another way.

Causing a polarisation of the membrane?

Yes. If that was the case, you could swap a positive proton for another positive ion like a sodium or a potassium, and you could swap a negative electron for another negative charge like a Cl- or Br-.

An uptake by a kind of anionic or cationic trading?

Yes, that's right. That was the hypothesis that I developed in the 1950s.

At that time work was being done in Kreb's laboratory in Sheffield by a man called R.E. Davies on the way hydrochloric acid is secreted by the gastric mucosa (in your stomach, my stomach, a frog's stomach, everybody's stomach). Davies and I got together in the late 1940s and decided that the hydrochloric acid secretion was possibly another facet of charge separation. There the proton was going straight off and the negative charge joining it was in the form of a chloride minus. That was a pretty exciting idea.

Now coming back to later in the 1950s: While I was at UCLA, I pointed out that possibly this charge separation had another function; it might be what activated the phosphate to go on to the ATP. So the charge separation was the basis both of ion movement and the formation of this energy-carrying substance, the ATP. I published this in 1960, and as far as I am able to judge, I think this was an original suggestion: that the charge separation could be used either for ion movement or phosphorylation.

Was the polarisation controlling which things could move?

That's right.

You were getting close to the cytochromes, in a way.

Yes, indeed. The separation was taking place at the cytochromes in the respiration story. The next step was in 1961 when Mitchell, the great star of this part of the game, came up with what he called the chemiosmotic hypothesis. This hypothesis, which he arrived at in parallel with my thinking and perhaps with some influence from me, made the point that this charge separation was likely to be both the active transport system and the ATP forming system. That was a great leap forward because he recognised something that I didn't have enough nous to see: if you reversed ATP and broke it down, you could get a charge separation back from it (that is, a positive and negative charge) across a membrane. That was where Mitchell and I had a very high degree of agreement starting in 1961 when we exchanged correspondence for the first time. He was then in Edinburgh.

Another influence in my thinking, which I haven't mentioned in sequence, was the great Swedish plant physiologist, Lundegårdh. The work that he was doing through the 1950s was very similar to what I was doing, and we exchanged information and were helpful to each other. I saw him in 1948 and then again in 1956. He had invited me to go and work with him in the late 1940s, but CSIRO thought I'd had enough leave and overseas trips and should stay in Australia.

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Overseas studies

Could I just ask about those overseas trips.

Yes, I had some wonderful opportunities to travel. In those days CSIRO was aware that we could easily be isolated in Australia, so they gave us the opportunity to travel for six or nine months to other parts of the world. In 1948 I went by ship to South Africa and travelled overland to see the fruit work in South Africa in particular. Then I went on to England where I saw plant physiologists and fruit storage, and got the chance to see Briggs again.

You stayed on and worked with him a bit?

Yes, I had about five months, a lot of it in Cambridge, and we produced a paper together. That was when I met Lundegårdh for the first time. Lundegårdh was not only a great scientist but an unusual man. He had enough money to build himself a private laboratory – which he did, adjoining his quite luxurious house some distance from his apartment in Uppsala. Incidentally, he was married to the daughter of Bruno Liljefors, a very famous Swedish painter. The Lundegårdh house and laboratory were hung with Liljefors' paintings, lovely things like a hare in the snow. In his scientific work, Lundegårdh was one of the greatest plant physiologists of the century, and he and I had similar ideas about the charge separation phenomenon and the quantitative relations between respiration and ion movement.

And you also travelled to the United States?

Yes, the opportunities to go to laboratories in America were splendid. My first contact on arriving in America was with David Goddard, who was a plant physiologist at Rochester, but he soon moved to the University of Pennsylvania in Philadelphia. He was very influential, and very good to me personally. He made contacts with the other plant physiologists in America, and told me whom I should go and see. So I met Kenneth Thimann, then at Harvard; Folke Skoog, who was at Wisconsin; and then across to the West Coast, which was the real stronghold of plant physiology in those days, with the Hoagland Laboratory. James Bonner was at the California Institute of Technology, and he was doing the remarkable work that took him into the biochemistry of plants. Also there was Fritz Went, the growth substances man. That was a great broadening experience that led subsequently to some collaborative work.

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Promoting scientific research

What about the CSIRO work in the late 1950s?

I mentioned that I went to UCLA in 1958-59. I came back because I was asked to join the Executive of CSIRO. CSIRO had about 4000 employees, and the Chairman, Sir Ian Clunies Ross, had died in office. The new Chairman, Sir Frederick White, asked me to join the Executive, which was a four-man body in those days.

I had misgivings about doing that, because I didn't think a full-time administrative job for the rest of my life was what I should do. First, I was still interested in research. And second, you do well at administrative jobs if you take on somebody else's problem and make it yours while you solve it. I found I could do it, but it took a great deal out of me and I didn't know whether I'd want to keep doing it for very long. So I took the administrative job in the CSIRO for up to three years, but in the event, I left after about two and half years. The Chairman didn't want me to leave, but he agreed that it would probably be better for me to take the next step, which was to become a Professor of Botany at Adelaide University. So that brings us to 1962.

You were better off in Adelaide?

Yes, it was like coming home – doing more plant physiology in a proper way, and I got the opportunity to make a couple more appointments. One of them was a man who had been working with Briggs, Professor Michael Pitman, who later became Chief Scientist in the Prime Minister's department in Canberra. The other was a former student of mine from Sydney who had been in America, Dr J.T. Wiskich. I brought him back in plant biochemistry, and he is still in the Adelaide department as a reader. So we had a strong physiology group there.

Did you still have a chance to do research at this stage?

Yes, I did, especially between 1962 and 1965. Then something happened that took away all my research time. The Australian government decided to do something it had never done before – namely, give individual grants to research workers or teams in universities. The minister in charge was Mr Gorton, who later became Prime Minister. At that time he was Minister Assisting the Prime Minister in matters relating to education and research. Some people, including the Chairman of the Universities Commission, thought that the money should be given in block grants to the universities. Others thought that there should be a system whereby the best work was supported; let it be competitive.

Mr Gorton sent for me from Adelaide to come and see him in Canberra, and I remember that when I walked into his office he was riffling some papers on his desk. He looked up and said, 'Oh, hello. Thank you for coming over. I wanted to ask you to be Chairman of a research grants committee that I'm setting up. It will be a bloody awful job and I wouldn't advise you to take it, but I'd be tremendously grateful if you would.' Who could resist such an offer?

So there you were, dealing with research grant applications.

The money was already available and there was a good deal of unhappiness in the universities because it hadn't been distributed at all. It had been delayed so long, we agreed that we should get going as fast as possible. Gorton had the idea that I should be able to do it with a committee of about a dozen people and, in fact, we started with ten. Those ten were specialists in different fields, but the system was based on confidential reports in writing from referees both in this country and elsewhere: 'This is Joe Blow. Here is what he is doing. He will be known to you through his research papers. How do you rate this man?' We brought in that system and distributed the available money. There were some disappointed people, but there were no rows. The system that I set up and operated as chairman for four years went very well and hardly changed in the next 25 years. It's changed now because the whole exercise has got rather bigger.

It was an important development that lasted for a quarter of a century.

Well, it was important to me; I felt it was something that was worth doing to help science in the community. That is probably why someone recommended me for a CMG [Companion of the Order of St Michael and St George], which I got at the end of that period. But it meant my research time was shot to pieces.

After the Adelaide professorship and your chairmanship of the research grants committee, you got an invitation to come back to Canberra to the Australian National University?

Yes, that's right, I became Master at University House. The mastership is like an Oxford or Cambridge mastership. The Master has a senior position in the university, is expected to be scholarly or scientific in his pursuits, has time to do so (or did in my time), and has some research assistance. I set up experimental work in the Research School of Biological Sciences and I had an assistant. I enjoyed being Master of University House because it was even more academic then than it is now. Indeed, we wore gowns to high table and had dinner in hall every night of the week in those days. My wife and I always stay there if we're in Canberra overnight.

It's a bit more commercial now, I suspect.

Yes, it has to be. In those days it was using some of the grants provided by the government. These days that's no longer possible.

Was your research a success in that time, because you would have been quite busy running University House.

Well, yes and no. The work was going quite well, but it wasn't University House that interfered with my time. Very soon after I came to Canberra, the President of the Academy of Science, Dr David Martyn, died in office. I seem to be fated to succeed people who had died in office.

You were captured again!

I was asked to be President of the Academy of Science. That, of course, is an honorary part-time job which becomes all-absorbing. The next four years, the four years of my tenure as President of the Academy, were at the expense of my research. I was able to do very little.

But you were busy for the Academy.

Yes, I did a lot of things. I suppose the most exciting, indeed the most straining, was when we entered a protest through the Australian government against the French atmospheric testing of nuclear weapons and suggested that there was a circulation of radioactive material. In 1973 the French asked if they could send four scientists to come and discuss this with representatives of the Academy. The discussion lasted over three days, during which time we agreed on all the technical details; we had a very happy atmosphere of discussion with our bright French colleagues.

We disagreed in the end because, while they didn't quarrel with the figures, they thought that the amount of radioactivity released into the atmosphere was negligible and likely to have negligible effects. We said that there was no way of knowing when a given dose of radioactivity was negligible, and we thought the prudent view was to stop atmospheric testing. That was a very interesting experience. It was run, I think, by the Department of External Affairs, and everything was done with strict protocol. Our French colleagues were such good chaps that it was an enjoyable experience, even though we differed in the conclusion. There were so many protests about atmospheric testing that the French stopped it soon after that. We may have made a very small contribution to that.

During your career in Canberra you moved to a new job.

Yes, I was appointed head of the Research School of Biological Sciences. I was there until retirement.

Where you facilitated a good many developments, rather than doing a lot of bench research, I believe.

Yes, that's true. I was asked by one of the great vice-chancellors, Sir John Crawford, the same one who had asked me to come to University House. He sent a message, saying the directorship of the Research School of Biological Sciences was coming vacant. That was in 1971, and he asked whether I would like to consider that rather than continue at University House. He asked Frank Fenner to come and discuss it with me. I think Frank was then the Director of the John Curtin School.

I was in hospital at the time Frank came. I was a very keen horseman, and I have the dubious distinction of having broken two legs on separate horses in one year – which was a bad thing to do, especially with the sort of break that requires some eight weeks of traction. But while I was in this state, the matter was discussed. I decided I would like to get back to a greater involvement in science and lesser involvement in administrative details which, though not enormous, were still significant in University House. So I accepted.

You didn't take long to decide I guess. Frank Fenner is terribly persuasive.

Yes, that's right. So that was how I came back to the Research School of Biological Sciences. I was still President of the Academy, which meant that I didn't start with a lot of time for research. I enjoyed the association with the research school people, an absolutely first-rate lot.

Even when I ceased to be President, yet another of these part-time jobs came along. This time it was the Prime Minister of Australia, Malcolm Fraser. I'd known him for some time, ever since CSIRO days when he'd come to visit us as a politician to see if we were doing our job properly. He wanted to set up something that the Academy had recommended: a science and technology council. The idea was to bring together ten or twelve people – bureaucrats from departments that had scientific activities, businessmen who had scientific or technological interest, and academics. Under Malcolm Fraser's persuasion I consented to be Deputy Chairman, and a former colleague from Adelaide University, Sir Geoffrey Badger, was Chairman, and that was no sinecure. I think my contributions through that (which ran into my period of retirement) might possibly have been the reason that I got recommended for the honour of Companion of the Order of Australia.

Well, you certainly put a lot into that job. We're into the last minute of the interview, Sir Rutherford, but there's a lot that you haven't talked about. I hope that we can meet again for a further interview.

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