Professor Jenny Graves, geneticist

Professor Jenny Graves

Professor Jenny Graves received a BSc Hons from the University of Adelaide in 1964 and an MSc in 1967. Her research for these degrees involved the inactivation of the X chromosome in marsupials. She then received a Fulbright Travel Grant to go to the University of California, Berkeley to work with Professor Dan Mazia. She received a PhD in 1971 for her work on the control of DNA synthesis. In 1971, Graves returned to Australia as a lecturer in genetics at La Trobe University. Her research interests focused again on marsupials and she became involved in gene mapping. Her current research involves investigating the organisation, function and evolution of mammalian sex chromosomes and sex determining genes. She is also interested in comparative genome mapping. Graves became professor of genetics at La Trobe in 1991, and became a Fellow of the Australian Academy of Science in 1999. In 2001 she took a position at the Research School of Biological Sciences, Australian National University as head of the Comparative Genomics Research Unit.


Interviewed by Professor Roger Short, 2000.

Contents


A biochemical geneticist career coming to fruition

I am speaking to Jennifer A Marshall Graves, of La Trobe University. Jenny, what does the A stand for?

Ann, my mother's name. And Marshall is my father's surname.

How long have you been at La Trobe University?

Since I was appointed to the Department of Genetics in its infancy, back in 1971. I was a very junior lecturer – I hadn't yet finished my PhD, let alone done a post-doc. It's been a very exciting 29 years.

You became a professor in 1991, and now you have moved into the Department of Biochemistry. Is this going to make you a biochemical geneticist?

I guess I've always been a biochemical geneticist, although I have confessed to the chairman of the department that I have never done very well in biochemistry and I've never really taken biochemistry courses. But biochemistry is increasingly taking over some of the genetic territory, like genomics – the physical structure of the genome and of chromosomes, and structure of genes – which I have been heavily involved in.

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The planting of the genetics seed

It would be fascinating to know what early influences channelled you into your eventual direction.

I've often wondered, as a matter of fact. Both my parents are scientists. My father was the head of the Soils Physics Subdivision at CSIRO. But I don't actually think that was what turned me to science. Certainly at school I never did much in science – the prizes I won were all art and creative literature. I wasn't being pushed towards science, but my family's example did make me realise that it was a viable option, and ordinary people were scientists.

Did you ever resent the amount of time that your parents devoted to their science as opposed to family? Did you see science as being in competition with family life?

Not at all. It never occurred to me. My father was sometimes in his own world. It didn't matter if you tiptoed around, because he took no notice anyway. My mother was working part-time when I was young. In fact, it really surprised me when I discovered that my friends' parents weren't scientists, they didn't work, they didn't have this other life. I just grew up assuming that everybody had their own life and their own interests which they fitted around their family and their relationships. I have a photograph of my father at an international congress – in Moscow, I think – when he was the Chairman of the Soil Science Society. I am particularly fond of that, because it looks like he is just about to open his mouth and welcome us.

As to choosing a career for myself, I remember in primary school they always asked you, 'What do you want to be when you grow up?' All the other little girls wanted to be air hostesses or nurses, but I knew I didn't want to be an air hostess. When I went home and asked my Mum and Dad what I should be, they said, 'Well, you're good at maths and art: maybe architect.' Once I found out how to spell it, for the next 10 years I put down 'Architect'. Then, when I topped the state in geography, I thought maybe I'd better do something geographical, and for the next three years I put down 'Meteorologist'. But it meant nothing at all; it was just nice to have something to write down, because it means that people treat you differently: 'Jenny really knows what she wants to do.' Jenny didn't have any idea, though – in fact, not till my final year at high school, when I had a very wonderful biology teacher. It was the first time I'd been exposed to biology, and I hated it – it was boring, a whole lot of stuff to learn – until the minute we got to genetics. All of a sudden I thought, 'Wow! This is easy, interesting, fun. I think that's what I want to do.' So when I went to uni the next year, I chose courses heading towards genetics.

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Growing towards the light: Honours work on gene structure and regulation

You once amazed me by saying that at first you actively disliked evolution.

Well, I had very strong opinions on what was fun and easy and what was difficult and hard. Evolution was definitely in the latter category – it was all formulae; the ideas were in books by R A Fisher, which I found a little bit impenetrable at the time – whereas physiological genetics, DNA, Watson and Crick, and messenger RNA were all new and topical and exciting. I decided, 'Gene structure and regulation is the direction I'm going into. I'll leave this evolution stuff well and truly alone.'

R A Fisher had a great Adelaide connection. Of all your teachers as an undergraduate at the University of Adelaide, who inspired you most?

Peter Martin's first-year lectures in biology were fascinating, wonderful. They sucked me right in. And as I became more senior in the genetics department, even though it was a very small department I had a number of very gifted teachers.

For my Honours I ended up – again completely accidentally – working with David Hayman on marsupials. The alternative was midges, and I came over to Melbourne to investigate working with them, but then I discovered that where midges grow is the sewage farm! I went off with Jon Martin to the sewage farm, and I decided I would rather work on mammals with David Hayman. He was a marvellous teacher and I got a great deal from his lectures and then also from the project I did in his lab. Peter Martin, who had always worked closely with David Hayman, was a co-supervisor in my Honours year, having come back from London with a terribly trendy new technique called autoradiography. So that was what I was going to be doing in my Honours.

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Roots spreading in unconventional ways: a Berkeley PhD

With a new degree hot in your pocket, you decided to go to Berkeley. How did you fund that?

The decision and the funding just happened. I can take very little credit for making a positive choice, except that I had actually been to Berkeley with my parents, who themselves got their higher degrees there. In fact, they more or less met at Berkeley. They had been there when I was a sophomore, and I had taken classes there – one from the wonderful Dan Mazia, one of my great heroes. He was a very famous cell biologist of the '60s, '70s and '80s, being regarded as a great god. He had a famous course which he called 'Physicochemical Biology' – today we would call it cell biology, but there was no such thing in those days. When I took that course, I felt for the first time, 'This is something I can really do.'

After I finished my Honours, having done better than I expected, I was offered a scholarship to stay in Australia. But I thought, 'No, I've had enough of being here and doing this. I want adventure.' So I said the first thing that came into my head: 'Oh, I thought I'd go and work with Dan Mazia.' And then, of course, I had to do it, which was simply terrifying. But he was very helpful and welcoming, and became my major professor at Berkeley. In applying to go to Berkeley I didn't think about the money or anything, but I got a Fulbright Travel Grant which helped out with the travel. And in fact I had a traineeship once I got there, a huge sum of $4000 a year – of which I paid half in fees, so I don't quite know how I lived there.

Had you ever felt, in your Adelaide setting, that it was very unconventional for a woman to think of a career in science? Or didn't it matter?

It wasn't the done thing. In fact, I was very conventional and I wasn't intending to do it. I wandered into a significant career by sheer accident; it certainly wasn't the first thing on my mind. I just kept wandering along and when there was a choice in the pathway I decided right or left more or less at random. The winds blew me towards subjects which I enjoyed and which ended up being on the road to something, but that's not really where I was at. I didn't actually buck the system at all until it came time to decide what to do after my Honours. That's when I decided – again absolutely on the spur of the moment, for the worst possible reasons – that I would go off to Berkeley and do a PhD there. But my mother had been a bucker of tradition. She had gone off to Toronto in the '30s to do her Masters, and that was definitely not what young ladies did in those days.

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Birdsong: adventures into marriage and a family

I guess one of the most exciting events for you at the University of California was meeting John. Can you tell us a bit about that?

I didn't meet him the first time I was there, though the funny thing is that he also took Mazia's class in Physicochemical Biology. But when I went back as a PhD student, he was also a PhD student in the same department. The story goes that we met singing on the same stage, but that isn't quite true. We actually met through Scientific French, which I regret to say we were both terribly bad at. Then the department put on a Christmas party for which one of the graduate students had written an entire opera,  Nucleoside Story. That was West Side Story converted to the story of the Biochemistry and the Molecular Biology Departments, who were at perpetual war. I was chosen to sing Maria from the Molecular Biology Department, and John was chosen to sing Tony. So we sang such song hits as 'There's a lab for us' and 'Urea'. It had a happy ending, because we ended up married and we've been married for 34 years now.

It was also an introduction to singing. I had never done any formal singing – I couldn't even sing from a score when I met John. But he had sung in a lot of very erudite little groups and so he urged me to try out for a group there. I can't imagine how I ever passed the audition, except that they were running rather late and they gave everybody the same score. It was a Dowland ballad which I had never seen in my life, but after three goes through with other candidates I knew it quite well! After getting into that group I had some wonderful experiences, singing with the university chorus and little choruses – I sang with Seiji Ozawa in San Francisco Opera House and with Stravinsky. And back here I sang with the Tudor Choristers for many years.

How did you persuade John to come to Victoria?

That was easy. He was very adventurous. He had planned to go to Europe that summer we met, but I was planning to come back to Australia because my sister was being married, so he thought he might as well come along. Then, since he was coming to Australia anyway, we thought we might as well get married too. My parents were quite surprised when we told them. Then, when I had finished my degree, I had to return to Australia because of the Fulbright grant conditions, so he came back with me. In fact, he loved Australia. He was born to be an Australian, I think.

How long did it take John to find his new career in your Eltham vineyard? He has been spectacularly successful at it.

His family has always been very interested in growing grapes and wine, and his young brother has set up a winery which is brilliantly successful over in California. John always felt that he wanted to do something too, at least at an amateur level. That has been building up over the years and they are now commercial, though small. But he has other careers too, and local politics has also been very important to him.

I like your photograph of him as Mayor of Eltham, with his mayoral chain. How long was he Mayor for?

For a year. He was the last Mayor of Eltham, which sounds like some kind of a novel! It was certainly quite a new experience for me to be Mayoress for a year, having to say the right things and so forth. It was good practice in getting out of the world of genetics for a while.

And you have two daughters. What are they doing now?

Erica is doing Arts/Science at Melbourne University, and Alison has just graduated with a Masters in Epidemiology from Berkeley and is now doing a PhD there.

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Looking to the local fauna: X inactivation in marsupials

Anyone who thinks of you in science, thinks of you in a marsupial context. Having had a marsupial start with David Hayman, when you came back from Berkeley with your PhD behind you how did you get involved in marsupials again?

Again that was completely by accident. When I went over to Berkeley I decided I was going to be a 'proper' molecular biologist and work on bacteriophages. But I had already been bitten by the bug of 'Aren't mammals fascinating?' The work I had done with David Hayman was about X chromosome inactivation – one X chromosome in females is actually genetically switched off – and that was a fascinating saga. Another of my heroes, Mary Lyon, was the one who put forward the hypothesis in the early '60s, and I was the one who tested it in marsupials. When I discovered that there is an inactive X in marsupials, I assumed it was just the same. It turns out that nothing is quite the same in marsupials.

When I came back to La Trobe University, Des Cooper – an old friend of mine from Adelaide – was a senior lecturer here. He was very keen to continue looking at X inactivation in marsupials, and we had to know what genes were on the X chromosome. He said, 'Well, you've had all this experience fusing cells, and that's one way we can map genes. How about doing some work on marsupials?' I remember to my chagrin and embarrassment pulling myself up to my full height and explaining that I was not going to be 'one of those Australians who end up working on “the local fauna”.' Only later did I see that there were good reasons for wanting to work on the local fauna. Just to keep Des quiet I made him a few hybrids, and that turned out to be so interesting that I continued some gene mapping work with marsupials. Gradually the realisation dawned that we had a goldmine there, that marsupials do things differently from placental mammals and that very often, if you compare those systems, you can figure out from how the two systems differ, what the ancestral system was like. So because of my interest in the regulation of genes and X chromosome activation, that became a very powerful way to look for variants.

When I first started, some people – Jim Peacock, particularly – said, 'They're going to be too different. You'll never find out anything.' When the first three genes I mapped were all exactly the same as in the mouse and the human, people started to say, 'They're all going to be the same so you won't find out anything.' But in fact they're just right: marsupials are just far enough distant from mouse and man to be interesting and to provide us with variation, but they're close enough to share the same control systems. Virtually everything I've done from then on has used comparisons between different groups of mammals, and most of it has turned out to be wonderfully interesting. I suppose I should have predicted that, but in the beginning I hadn't seen the possibilities.

It's interesting how long it has taken the rest of the world to wake up to the intellectual excitement of Australia's isolation for 100 million years.

That's quite true. When I first started getting interested in travelling and talking about my work – which took me some time because I had young children – if I mentioned a wallaroo, the audience would all fall off their chairs laughing. I became a bit of a comic turn, I think. I'd show lots of pictures of marsupials, and they'd all think, 'Oh, this is such cute stuff.' I don't think they took it seriously until the saga of David Page's ZFY gene: it pulled people up with a jolt to realise that marsupials could be a test of what should be truly conserved.

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The cherry tree: tracing career developments

I remember that when you gave an address here at La Trobe, after you had been elected to the Australian Academy of Science, you showed us a lovely 'cherry tree', with all your research grants as leaves and all the published papers as cherries. That was a great way of summarising your scientific career. I'd love to go through it with you.

That was something I had done many years ago, because I wanted for my own purposes to see how my career had developed. I carried it on and the little tree grew and grew. I found it quite fascinating to look at the accidents that attend any career and how you end up with different, interwoven pathways – you can't really tell, when you're down at the beginning of the tree, which bits are going to be successful and which ones aren't. Many years later you can look back and see that there were a number of branches that didn't lead anywhere, but other branches that started off rather weak and spindly started to get fatter and grow.

Can you take us through some of those branches and fruitings?

Yes, I'll do that. The tree shows where I was at in Adelaide, working on marsupial X inactivation. That led up to my time in Berkeley when I started working on the factors involved in DNA synthesis control – that was what I had been working on in marsupials, so I generalised it to working on animal cell hybrids and that's really what I was intending to spend my entire career on. But that branch fizzled out, and what took over was work which grew out of Des Cooper's comments that I ought to start working on marsupials, because this is where I started becoming very interested in Australian animals. And although that seemed like a teeny-weeny twig then, it grew into being almost the whole tree.

Cherries, leaves and branches: where the genetics work has led

Which was your biggest 'cherry', and which was your biggest 'leaf'?

One of my biggest cherries came with the sex determination work. That was a very important cherry, because that was our demonstration that the gene we thought was the sex determining gene actually wasn't. The tree has gold cherries, too, for the three books which I've edited so far. Tracing the branches up: some have lots of cherries and lots of leaves – that is, they were very well supported and very successful, with lots of publications. I also had little side-projects on irradiation and position effect variegation which then I never was able to 'leaf' very well. They were under-resourced and they generally dried up after a cherry or two.

Where the grant support was forthcoming was in the gene mapping work that I got into during the '80s. That came from my work with marsupials, and was very important because it led me to become part of the comparative mapping committee of the International Workshops on Human Gene Mapping. That was great, because here at last I had some people who appreciated what comparisons could do for you, and everybody was very keen to include marsupials in these comparisons because they are so distantly related. So everything has come off those branches, including work on genomics – comparing the whole genome. We take an array of genes from one species, usually human, and ask, 'Where are these pieces of the jigsaw puzzle in a marsupial? How have they been rearranged over the last 130 million years?' And that we have done in two ways: by mapping genes and also by the new technique of chromosome painting, which we really love because it's so very beautiful. That has been extremely successful, and we have a nice big fat leaf there – my biggest up till last year when a slightly bigger one 'grew' on the tree. The work it supported has also been very fruitful: we've had numbers of publications from it. And it has led to a lot of interaction.

This branch was the work I was doing on X chromosome inactivation. I did some rather important work during sabbaticals in Seattle and in San Francisco, but really I never was able to get support in Australia. So that part of the tree intertwines with the 'mapping' part and feeds off it. It is still going and I think it's going to take off and be a very important branch in the next few years. Hopefully, we will be able to isolate the important control gene from the marsupial X, and we'll be able to compare those as well. The 'sex determination' branch and the 'Y chromosome' branch have also come off this 'gene mapping in weird mammals' branch.

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Pruning or nourishment? The crucial role of research funding

With such a very fruitful tree, has anyone ever made the terrible statement to you that you might have been more productive if there had been a bit of judicious pruning?

Only me! In fact, I've had no interference directly from anyone, ever, in my entire career. Nobody in the university has made any suggestion, helpful or unhelpful. But of course you are very much at the mercy of funding agencies – a lot of the directions I have gone in have been driven by funding decisions. I would very much have liked to pursue some things, but it was clear that the funding was simply not there.

These days we are being told all the time that we have to start pursuing more applied research which is of interest to companies, who will therefore lavishly fund us. And in the current funding climate, that is probably very realistic. So I have given a bit of thought to the direction in which I might take my research, where it would be of some interest in its applications, and I have now put in for some small funding. But I don't want the tail to wag the dog, and I certainly wouldn't wish to be in a position where I was being constrained by having to have a certain result or even go in a certain direction.

I'm afraid the days when we can expect to be quite so free in our research are coming to a close, but I've been very lucky, in that I haven't been constrained. I think there would have been no way I could convince some company to invest in gene mapping in marsupials. I wouldn't myself have been able to mount a good argument. I can now, in retrospect, but there were so many surprising things that came out of it, from the discovery of the sex determining gene to all sorts of things that didn't work the same way in marsupials as they did in humans. We could never have predicted that.

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Putting on new foliage: the endless task of grant applications

This is the sort of personal question one never dares ask even one's closest friend, so forgive me: How many of your research grant applications have been successful?

Probably not that many more than the national average. Sometimes I'm very successful, and sometimes very unsuccessful. One year I lost four out of four grants. That was while I was in hospital for a year after neurosurgery, too, so I honestly thought that was going to be the end of my career. But then the next year I got five out of five. It's very up and down, which makes it just about impossible to do any long-term planning. Like many people, perhaps, I simply apply for a lot of grants. I'll spend probably two months over Christmas doing nothing but applying for grants. That is not very productive, in that it doesn't put cherries on the tree, but it does focus the thinking something wonderful. I could certainly wish for longer and bigger grants, because they would allow me to do more work and less grant writing, but that's the reality. Perhaps most people don't realise that. They think, 'Oh, you're always successful at grants.' Not true at all – maybe somewhere between a half and a quarter.

When you've been turned down for grants, but then you have been able to fund the work independently through other sources or somebody else has gone ahead and made a success of it, have you ever felt, 'I told you so!'?

Yes, of course I have. There's nothing better than the feeling, 'I told you I could do it, and I did it!' Certainly I have done that a few times in recent years. Frequently, the very thing you need to mount a successful application is the demonstration that you actually have gone some ways along and you do have some preliminary results. I have certainly made use of preliminary results. I'm often a little bit ahead of where I say I am at in the grant application, so I know I can do it.

I think we all do that. But you have touched on an interesting point, that in many ways adversity is the spur to success.

Well, a little adversity is fine, but maybe not a lot of adversity. The year I had no money and I was in hospital was definitely not a spur to my career. S

Overcoming a health setback

Would you tell us a little about your health problem? When we heard about it we all thought, 'Gosh, bye-bye Jenny.' To have made such a comeback is fantastic.

It was a big shock. I came back from a very successful trip to Los Angeles, got off the plane and went home, and in 20 minutes my balance just fell to pieces and I started to see double. So I knew I was in deep trouble. It turned out I had a haemangioma in the fourth ventricle, which was a good thing to have, but not where you keep your heartbeat and respiration – they call it 'tiger country'. When that was operated on, I was warned it would be 18 months before I was really back on my feet – and it was 18 months before I was 100 per cent better. I was absolutely grimly determined to stick with it, so I was back at work in three or four months. But I'd get to work and then have to turn around and go home again because I was so exhausted. It took a few years off my life, I think, but I was determined to get back into some kind of normal working frame of mind – very, very determined to get my career back.

We watched you in those months afterwards, having to hold onto a doorpost or something because you were still quite wobbly. It must have required great courage to persevere and put in those grant applications, and keep the lab running.

You don't get much choice! I had some wonderful people in the lab. But with all my grants gone, I couldn't fund the thirteen people in my lab, so a lot of people left the lab and I was in despair. The few people who were left were incredibly helpful to me. I could really not have done it without them.

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Nurturing the seedlings: a superb teacher

Let's talk a bit about the lab personalities. I will always think of you as a superb teacher. You have inspired a generation of graduate students who have gone on and done great things. What is the key to your great success as a teacher?

I've often wondered. I have had wonderful graduate students, although some of them didn't seem that wonderful when they came. I've often picked up students that for one reason or another have not been very successful – in the early part of my career, particularly, eagerness was what I was looking for. Even if somebody didn't have the 'best' first-class Honours, I invested a lot of time in them if I thought they were truly motivated and truly eager and interested in what they were doing. I suppose that's always what I've been looking for.

I love my lab. It's an absolutely wonderful place for me, full of enthusiasm and full of debate and interest. I have for a number of years had five or six or seven – and currently eight – postgraduate students, and I love 'em all! I never want them to leave, but of course they have to. I've been lucky, in that I have been able to follow their careers. I feel like I've got a network of ex-JennyTech people all over the world. I still have a great deal to do with a number of them and we collaborate a lot. It's a huge pleasure to me to see them succeed – like being Mother all over again, a mother of 20 PhD who are all doing interesting things.

At the end of last year, when I was elected to the Academy, a number of my old students who were in my lab many years ago – I think the earliest was from 1972 – came back to a dinner at which they wished me and my current crop well.

You've always done more than your fair share of undergraduate teaching. Do you find that enjoyable, or a bit of a grind?

Both, of course. I've just started my undergraduate subject in the last couple of weeks, so I'm very much into exactly how I feel. I always go through a terrible slump as it looms up closer and closer, because I'm involved in trying to get this done and that done, and send papers off, et cetera. I can see this great shadow arriving, and so I get very depressed. But I know that the minute I start teaching I'll feel really terrific about it. Now I know that will happen, it's easier to cope with.

I enjoy the exercise of putting things before a class, and in a way that they will not just understand but really be enthusiastic about. My course in third year is on human molecular genetics; that is a winner because they are always interested in it, so I thoroughly enjoy teaching it. But it is incredibly exhausting – not so much the lectures, but the lab classes. I've got four days of four hours a week. Yesterday I was talking for five hours solid, until I said, 'Enough! At this rate, I won't be able to say anything to Roger tomorrow.' I do enjoy teaching, but it is a huge drain of energy. And if I did less teaching, I would get a lot more research done. Can't do everything.

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Stresses and strengths: the importance of doing what matters

You must also have been a great role model to Australian women in science: you are an example that someone can make a career in science and reach the top, yet also be a mother, a wife and a successful home person – who gives fabulous parties, if I may say so.

Part of that is my husband's wine!

Do you think you have drawn more female students into research than a male chairman of a department might have done?

Probably not. I haven't had a lot of female students, probably no more than anybody else. I'm often asked by women who are considering science as a career, 'How do you do it?' and my answer is, 'H-a-a-h, you work incredibly hard,' because that is still the case. I'm frankly not sure that any of the committees I've ever been on about women in research and so forth are terribly effective. Things haven't changed all that much. You just are doing two jobs: you work incredibly hard, particularly when you have young children. People can help you – John was a big help – but essentially there is nobody who can do it except you. You just have to put in the energy and the hours.

I must say that when I first had children, it did amazingly focus the mind. For perhaps the first time I started to really value my time. Before that I would come to work and spend the day pleasantly, humming a little tune, looking at my cells through the microscope, maybe without anything specific in mind. That all ceased in a minute. Right away I realised I couldn't keep on going unless I called my shots. It was very good for me to have to go through that exercise of valuing myself and my time a lot more highly.

What have you found to be the greatest stresses in your scientific career? And what have you found the best way to relax?

I think I'm not a very stress-y person. I am often asked, 'How can you be so relaxed, when the sky is falling, the sky is falling!' I guess I know that eventually I can cope with most things that happen. I have no trouble relaxing. A lot of things that I do have been extremely relaxing: singing, just being outdoors, my home life. So that's never been a real problem.

The biggest stresses were, and still are, competing claims on my time. Being a university scientist has always been an absolutely impossible job, but it has got more and more impossible in the things you're expected to do. I work 70-odd hours a week, as I think most of us do. That doesn't bother me. What bothers and stresses me is just the increasing amount of work that doesn't seem to have any purpose, but that you have to do or something dreadful will happen. The demands become more and more extreme – endless reporting and accounting and counting up papers and dividing them by two, and rubbish like that. Things that I can see purpose in, I don't mind putting a lot of effort into. But wasting my time is very stressful.

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An orchard-full of stimulating ideas

So, Jenny, a final question: What would you most like to be remembered for?

For ideas. When I look at some of my heroes and the contributions they've made, Susumo Ohno stands out as somebody who excited me as a 17- or 18-year-old reading some of his first efforts – not because of the facts but because of the ideas that were so iconoclastic then, and still are with us. They are still the framework for a lot of the questions I'm pursuing. Many are wrong but they were all interesting. I remember knocking on his door at the City of Hope a number of years ago to announce to him that we had just broken Ohno's Law. This was the first time that anybody had shown that genes on the X chromosome in one species weren't on the X chromosome in others – marsupials. Susumo Ohno has been a great hero throughout all my professional life.

More and more, as I gain in confidence, I like to think that some of my ideas may be more important than the actual data that we produce, although that has been surprising and interesting too. I'd like to be remembered for having put on the table some quite shocking ideas: first of all that the gene we all knew and loved was the wrong gene, and then that the right gene doesn't work the way we think it should, and the Spermatogenesis gene actually has a friend on the X chromosome. The evolution of the sex chromosomes is quite different from what we thought it was.

I get a big giggle every time my research points me in the direction of realising that maybe things are not the way we thought they were, and I've got a lot of pleasure out of developing some of these ideas in reviews and papers. Currently I am thoroughly enjoying writing a book which I hope will be a way of presenting ideas in the way that Susumo Ohno presented his. Maybe some 17-year-olds will read it and think, 'Oh, wow! This is fun. I think I want to do that.'

Jenny Graves, you've produced a lot of cherries, and many of us are going to be savouring those for the rest of our lives. Thank you very much indeed for sharing all these inspiring thoughts with us.

Thank you, Roger. It's been a pleasure talking to you, as ever.

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