Why, to my wife’s dismay, I made a late academic career move

I decided early on in my undergraduate career that I wanted to be a university teacher, so I knew that I would have to do a PhD, which is what I did indeed go on to do.  My first degree, Agricultural Zoology (essentially applied entomology and parasitology), had engendered a love of the applied and my PhD and subsequent post-docs echoed this, I worked on cereal aphids and ways in which to control them without recourse to pesticides. I had noticed as an undergraduate that although my degree was applied, the staff teaching us, had not actually worked outside universities.  I felt that if I was going to be a successful applied academic I should have some experience outwith the university system and in 1982 found myself working as a research and advisory entomologist for the UK Forestry Commission based at their Northern Research Station just outside Edinburgh (Roslin). My original plan had been to stay there for about five years and then move into a Lectureship. Unfortunately, 1987 coincided with a dearth of positions within academia and it was not until

Senior Scientific Officer, Entomologist with the Forestry Commission Research

1992 that I made the transition, when I was appointed as Lecturer in Integrated Pest Management* at Imperial College, London, based at the Silwood Park campus.

Newly appointed Lecturer in Integrated Pest Management 1992

At the age of 37, and armed with 50 publications, six children and a spouse in tow, I think this probably counted as a mid-career move.  It was a great time to move to Silwood. The Centre for Population Biology, headed up by John Lawton was in full swing, with future luminaries such as Chris Thomas, Hefin Jones, Brad Hawkins, Bill Kunin, Andy Hector, Mike Hochberg, Suzanne Koptur and Shahid Naeem in post.  In what was then the Pure & Applied Biology Department, I was one of eighteen entomologists, pure heaven. Silwood Park had a vibrant coffee culture, in the morning over at the Refectory (Paddy’s), where bacon and egg butties (rolls)) were consumed in huge numbers and in the afternoon, tea was served by Pearl over at the Conservatory at the rear of the Main House.  In the summer, we would sprawl on the grass outside, exchanging ideas and discussing our research plans.  There were no silos, and there was a significant amount of inter-disciplinary mixing.  The ideal environment for productive and innovative research and I was content to imagine myself in situ until retirement beckoned.

Unbeknownst to me there was something rotten in the offing.  Back in 1986 the University Grants Committee assessed the quality of research being produced by UK universities to help guide them in allocating research funding. The Research Selectivity Exercise and the Research Assessment Exercise followed this in 1989 in 1992. The criteria used to assess university research standing, was, largely based on the impact factor of the journals that staff published in and the amount of research income won from research councils. The effect across those universities in the top tier was insidious but largely predictable. Despite over 50% of income being accrued from teaching, recruitment of staff became increasingly based on publications, leading in the case of the Biology Department at Imperial, to an influx of staff with publications in Science, Nature, PNAS and similarly high impact journals, much to the frustration of our Director of Teaching.  Effectively, every time a whole organism biologist retired or left, they were replaced by a molecular biologist.

Things got even worse when in 2001, Sir Richard Sykes was appointed as Rector (Vice Chancellor).  Coming from industry and with a reputation for asset stripping, the collegiate nature of Imperial College came under attack. Staff, who had, until then, been valued for their departmental contribution in terms of administrative roles and contribution to teaching now felt under threat and if unable to up their research profile, encouraged to seek early retirement, or in some cases, as when Imperial acquired Wye College, which was subsequently closed, made redundant. Despite having a large research group, albeit largely made up of PhD students my teaching load continued to increase.  I even ended up teaching forestry and plant biology, when the plant scientists were culled. Another example of how research trumped teaching was In 2007, when despite me pointing out that the youngest whole organism biologist was 52 (me), and the oldest molecular biologist 52, our Head of Department decided that we needed more systems biologists.

In terms of career progression, I had in the early days, done quite well having been promoted to Senior Lecturer in 1996 and Reader in 2002. By then, looking at Athene Donald’s list I see that I was pretty much doing a professorial role, serving on external committees, validating degrees for other universities and acting as an external examiner. I was also appointed as Editor-in-Chief of Insect Conservation and Diversity, a new journal for the Royal Entomological Society. My administrative duties had also continued to increase.  It was no wonder that my beard was getting greyer! I was however still preparing my own talks, although I will confess that a lot of my data analysis was being passed on to members of the group, duly acknowledged of course. I am extremely grateful that I have always had a loyal and very supportive research group, without their help life would have been impossible.  My thanks to you all (if any of you are reading this). Promotion to full Professor was by departmental or self-nomination and successful interview performance in front of a panel composed of the Principal of the School of Life Sciences and Deans from across the university. If you wanted to be nominated for promotion one had to submit a lengthy cv to your Head of Department and this was considered by the other Professors within the Department to see if you were worthy of being put forward. In 2010 I had two post-docs and eleven PhD students and 136 publications, so felt that I was ripe for promotion to full professor. To my disappointment, the Department thought otherwise and my name was not put forward. I shrugged my shoulders and decided that there was no point in trying again, especially as by 2011, I only had one post-doc so that year I didn’t bother putting my cv in despite being pressed by the Department to do so. The departmental panel however, to my annoyance, considered my cv in absentia and pretty much forced me to submit even though I pointed out that it was going to be a waste of time as my research group was now smaller. Nevertheless my cv was submitted to the university and I was duly called for interview.

Accompanied by my Head of Department I arrived in good time for my scheduled 45-minute interview.  The panel finally called me in twenty minutes late. The Principal of the School of Life Sciences, an immunologist of some repute, began by saying “Hello Simon, can you tell me what you have been doing since you were last promoted, if you can remember that far back”. I looked at the HR person in shock, expecting some sort of response but nope.  Anyway, I proceeded to tell the panel what I had been doing since 2002. At the end of my reply, her response was “So you count things”.  That immediately told me that I was not going to get promoted.  The questioning passed on to one of the Deans, who said, in an accusatory manner, “I see that you get a lot of prizes for teaching”. I explained that I liked students and felt that they deserved the best experience I could give them. That went down like a lead balloon. The second Dean said “I see that you don’t give very many plenary lectures”.  I replied that I received many invitations but that my teaching load made it difficult to accept them all. Next up was the third Dean, who remarked on the fact that I only had one post-doc and that my research group was mainly PhD students.  I replied that yes, this was the case but that for the last 18 years I had been providing the Department with a steady and substantial income.  Then back to the Principal, who asked me why I hadn’t published in Science and Nature recently. Those of you who are applied entomologists will know the answer to that one 🙂 So that was it, twenty five minutes and I was dismissed, the interview panel were back on schedule, and of course I was not promoted.  My Department, to give them their due, appealed against the decision, pointing out that that I was actually doing more teaching than anyone else in the Department including the Teaching Fellows and that my research income was above the Departmental average.  The Faculty response was that perhaps if I did more teaching they could consider me for promotion via a teaching route.  I laughed hollowly. That was pretty much the final straw, it was obvious that the Faculty of Life Sciences had no interest in my research. My Department on the other hand relied on me and the other whole organism biologists to run six MSc courses and to service several undergraduate modules**.

It became increasingly obvious that things could not carry on as they were. My teaching and administrative loads were becoming ridiculous.  There were only four entomologists left in the Department and the coffee culture had almost disappeared, most groups staying in their silos and only venturing out at lunchtimes. I was seriously considering early retirement although I was reluctant to do this, as I was sure that with my retirement the last entomology degree in the UK would quickly disappear.  Then Peter Mills, Deputy Vice-Chancellor of Harper Adams University, approached me and asked what it would take to get us surviving applied entomologists to move.  I told him and was reassured that this was all possible.  With some trepidation, I agreed and we moved despite some half-hearted attempts by the Department to get us to stay.  I don’t think that they actually believed that we would leave a world-class institute to move somewhere that they had never heard of. I have never regretted the move. It was great to be in an environment where our skills were appreciated and back in a collegiate and collaborative atmosphere with an excellent coffee culture and to cap it all, better research facilities than we had had at Silwood. I take some pleasure in pointing out that I moved from a research intensive institute to a more teaching focused one, but halved my teaching and administrative loads. As a result of having that extra thinking time I very soon won two research grants. Domestically it was a bit hard as my wife couldn’t find a similar job in Shropshire so we ended up as two-house family, commuting between each household on alternate weekends, but entomology in the UK at postgraduate and undergraduate level is now in safe hands with a number of newly appointed entomological staff.  Though as my wife points out, if I had taken early retirement we would now be safely ensconced in France with full residency and not being subjected to the horror that is BREXIT.

*the last time that my Department advertised so specifically for a post.  Subsequent job adverts were along the lines of “in the area of ecology, evolutionary biology…”

**I confess I was rather pleased when I heard that Imperial had to close those six MSc courses because of a dearth of staff able to teach on them, although I was sad that after almost a century, postgraduate entomology was no longer being taught at Imperial.

The Academic Work-Life Balance – Doing what you enjoy for as long as you can

I am very lucky.  Unlike many people, I have essentially been paid to do what I love for my whole life.  My job is my hobby, my life even. I get paid to study and talk about the natural world, insects in particular, and have done so for the past forty years.   How lucky can a person be?   That said, it hasn’t been 100% fun all the way.

As I enter semi-retirement (3 days a week) I thought I would be self-indulgent and reflective (navel gazing in other words) and share a few thoughts about my academic work-life balance past, present and future.

http://fineartamerica.com/featured/the-unbalanced-scales-stevn-dutton.html

 

As a PhD student the scales were very heavy on the research side.  Apart from some demonstrating in the labs and a few Maths tutorials (BIO101) it was reading, writing and research.  Albeit this involved weekend working, but as there was plenty of time doing the week to fit in games of squash (our lab had a very competitive squash ladder) between field and lab work, it was pretty much fun all the way.

The PhD and first job– research heavy, a fun time

My first ‘permanent’ job was with the Forestry Commission, where I was based at their Northern Research Station, just outside Edinburgh.  My first few years were almost idyllic, lots of field work in remote parts of Scotland, the ability to have PhD students, giving guest lectures at Edinburgh and Aberdeen Universities, and an official ‘side-project’ time allowance which allowed me to write papers on a diverse range of subjects not included in my job description, e.g. my foray into species-area relationships (Leather, 1985,1986,1990,1991).  By the end of my time there however, government policy had changed, and we, even as a research organisation, were very much ‘customer facing’ and freedom to do less applied research was very much restricted to our own time.

Early academic life – when grant writing had some rewards and didn’t seem to take up as much time

It was thus a huge relief when I joined Imperial College at their world famous, and at the time, very collegiate, Silwood Park campus.  I was able to have coffee with luminaries such as Mike Way, Mike Hassell, John Lawton, Stuart McNeill, Val Brown and Nigel Bell as well as to rub shoulders with up and coming stars such as Sharon Lawler, Lindsay Turnbull, Jeremy Fox,  Chris Thomas, Shahid Naeem, Mike Hochberg, Charles Godfray and many others.  I could research any topic I wanted to as long as I got funding (and I did) and my teaching load, if not as light as some within the department, was manageable and very enjoyable.

It starts to tip

 

Administration has never been my thing, but as I got more senior, more administrative stuff came my way, and in my last few years at Imperial College where I was the Postgraduate Tutor, a role combining pastoral care and regulatory matters, such a chairing all the MSc exam boards and monitoring PhD student progress.  Luckily, I was very ably helped by two fantastic people, Diana Anderson and Janet Phipps.  Without them my life would have been a misery and the paperwork in an awful mess, to put it mildly.   I also ended up on a lot of college committees as well as taking on a number of external roles; editing, refereeing, external examining etc.  At the same time, Imperial College, as a joint consequence of appointing Sir Richard Sykes as Rector and the Life Sciences Faculty adopting a largely publication metric-based approach to new appointments, started to replace retiring whole organism biologists and entomologists with molecular biologists and mathematical ecologists.  Not necessarily a bad thing if managed sympathetically, but they still expected the same course content to be delivered by the few remaining whole organism biologists.  To give you an idea, when I joined the Department in 1992 there were 18 entomologists, when I left there were three of us.

My teaching load soared, while the departmental average was 25 hours per year, my personal load was 384 hours and I was also having to run a research group! The collegiate atmosphere was also very much eroded as was the attitude toward students.  When I first started at Imperial as a Lecturer, only Senior Lecturers and above could “Process” at the graduation ceremony in the Albert Hall. By the time I left, Teaching Fellows were being asked if they would like to attend. The majority of Faculty saw no benefit to them in attending.  A sorry state of affairs as far as I was, and am concerned.  Seeing our graduates happy and smiling with their families is such a buzz; why would anyone want to miss that?  We also had a change in our Director of Undergraduate Studies (DUGS), our former DUGS ‘s philosophy was to give the students the best possible experience with the resources available.  Our new DUGS’s was completely different.  His opening address to the Faculty went along the lines of “I know you don’t like teaching…” (this upset quite a few of us who did and do enjoy teaching) and his underlying philosophy was, as far as I could make out, how can we make the students think they are getting a great experience without expending too much time on them.  I was very pleased to make my move to Harper Adams University in 2012* where collegiality and student provision were, and still are, very much more valued; all Faculty are expected to attend the student graduation event unless they have a very good excuse😊

The things I have disliked the most over my career are grant applications, over long committee meetings, unnecessarily complex paperwork, office politics and marking assignments and exams.  On the plus side have been my good colleagues, lecturing, field courses, research project supervision at all levels, the opportunities to do outreach, and the students who have made it all worthwhile.

With retirement comes the opportunity to dump most, if not all, the things I dislike, and to concentrate my efforts on those aspects of the job I love the most, teaching, outreach and writing.  In the main, I have had a great time as an academic, but in the present climate, I would think very hard about advising my PhD students to take up an appointment in a Research Intensive university in the UK, especially if the value their family life and their mental well-being.

Hoping to spend more time in France 😊  The biggest challenge will be developing the ability to say no.

 

 

References

Leather, S.R. (1985)  Does the bird cherry have its ‘fair share’ of insect pests ? An appraisal of the species-area relationships of the phytophagous insects associated with British Prunus species. Ecological Entomology 10, 43-56.

Leather, S.R. (1986)  Insect species richness of the British Rosaceae: the importance of host range, plant architecture, age of establishment, taxonomic isolation and species-area relationships. Journal of Animal Ecology 55, 841-860.

Leather, S.R. (1990)  The analysis of species-area relationships, with particular reference to macrolepidoptera on Rosaceae: how important is data-set quality ?. The Entomologist 109, 8-16.

Leather, S.R. (1991)  Feeding specialisation and host distribution of British and Finnish Prunus feeding macrolepidoptera. Oikos 60, 40-48.

*It may amuse my readers to know that I applied to Harper Adams (then an Agricultural College) for a Lectureship in Ecology in 1992, for which I was rejected.  The following week I was offered the job at Imperial 🙂

Red, green or gold? Autumn colours and aphid host choice

“The falling leaves
Drift by my window
The falling leaves
Of red and gold”

Red, green and gold, all on one tree

When Frank Sinatra sang Autumn Leaves he was almost certainly not thinking of aphids and I am pretty certain that the English lyricist, Johnny Mercer, who translated the words from the original French by Jacques Prévert wasn’t either 🙂

The colours we see in autumn are mainly due to two classes of pigment, the carotenoids (yellow-orange; think carrot) and the anthocyanins (red-purple).  Carotenoids are present in the leaves all year round but are masked by the green chlorophyll.  Chlorophyll breaks down in autumn, leaving the yellow carotenes visible.  The anthocyanins on the other hand are not formed until autumn (Sanger, 1971; Lee & Gould, 2002) and this mixture of pigments give us the colours that have inspired so many artists.

Autumn Leaves Georgia O’Keeffe (1924) Tate Modern

To many, autumn starts with the appearance of the first turning leaves, to me it is the arrival of gynoparae* of the bird cherry-oat aphid (Rhopalosiphum padi) on my bird cherry (Prunus padus) trees.

Bird cherry, Prunus padus, leaves on the turn.

Little did I know when I started my PhD in 1977 that almost thirty years later I would be part of a raging debate about the function of autumn colouration in woody plants. At the time I was interested in the colonisation patterns (or as I pretentiously termed it in my thesis ‘remigration’) of bird cherry aphids from their secondary grass and cereal host plants to their primary host bird cherry.  My study system was 30 bird cherry saplings divided between two cold frames in the Biology Compound at the University of East Anglia (Norwich).  Every day from the middle of August until leaf fall I checked every leaf of each tree, for gynoparae, males and oviparae, carefully noting the position of each leaf, its phenological stage and giving it a unique number. I repeated this in the autumns of 1978 and 1979.  The phenological stage was based on the leaf colour: green, mature; yellow, mature to senescent; red, senescent.  What I reported was that more gynoparae landed on green and yellow leaves than on red and that the gynoparae on green and yellow leaves survived for longer and produced more offspring (oviparae), than those on red leaves (Leather, 1981).   The gynoparae of the bird cherry aphid are quite special in that although as adults they do not feed (Leather, 1982), they do not land on bird cherry trees at random (Leather & Lehti, 1982), but choose trees that not only do their offspring (the oviparae) do better on, but that also favour those aphids hatching from eggs in the spring (Leather, 1986).  It should not have come as a surprise then, that when I analysed some of the data I had collected all those years ago, their preference for green and yellow leaves over red ones, is linked to how long those

Figure 1. Length of time leaves remained on tree after first colonisation by gynoparae of Rhopalosiphum padi (F = 30.1 df 2/77, P <0.001)

leaves have left to live (Figure 1). The timing of events at this time of year, has, of necessity, got to be very precise. The egg-laying females (oviparae) are unable to develop on mature bird cherry leaves (Leather & Dixon, 1981), but it seems that the bird cherry aphid has this under control, making its decisions about the timing of the production of autumn forms (morphs) sometime in August (Ward et al., 1984).  All very sensible as far as I was concerned and that was as far as I took things.  Subsequent work by Furuta (1986) supported this in that he showed that maple aphids settled on and reproduced on green-yellow and yellow-orange leaves but avoided red leaves which had shorter life spans.

Jump forward fifteen years or so, and in a paper, that at the time, had somehow passed me by, the late great Bill Hamilton and Sam Brown (Hamilton & Brown, 2001) hypothesised that trees with an intense autumn display, similarly to those brightly coloured animals that signal their distastefulness with yellows, blacks and reds, were signalling their unsuitability as a host plant to aphids.  Like the costs imposed on insects that sequester plant toxins to protect themselves against predators, the production of anthocyanins responsible for the red autumn colouration is expensive, especially when you consider that the leaves have only a short time left to live (Hoch et al., 2001).  In autumn, trees and woody shrubs are normally mobilising resources in the leaves and moving them back into themselves ready to be used again the following spring (Dixon, 1963). Ecologists and evolutionary biologists were thus keen to explain the phenomenon in terms of trade-offs, for example, fruit flags that advertise the position of fruits for those trees that rely on seed dispersal by vertebrates (Stiles, 1982) or as ultra-violet screens to prevent tissue damage (Merzlyak & Gittelson, 1995).  Hamilton & Brown felt that these hypotheses were either, in the case of the fruit flag, only applicable to trees with fruit present and, in the latter, untenable. Instead they advocated the ‘signalling hypothesis’ which was based on the premise that trees that suffer from a lot of aphids (attacked by more than one species rather than by large numbers of a single species), invest in greater levels of defence and in autumn advertise this using bright warning colours.   The premise being, that although it is metabolically expensive for the plants to produce these colours, it is worth the investment if they result in a reduction in aphid attack.

This hypothesis was not without its detractors. Others suggested, that far from avoiding red colours, aphids were attracted to yellow or green as an indicator of host nutrition (Wilkinson et al., (2002).  Holopainen & Peltonen (2002) also suggested that birch aphids use the onset of autumn colours to pick out those trees where nutrient retranslocation was happening, and thus with higher levels of soluble nitrogen in the leaves.  This was of course, what I was trying to confirm back when I was doing my PhD.  Conversely, supporters of the signalling hypothesis, argued that trees (birch again) that could ‘afford’ to produce bright autumn colours were fitter, so more resistant in general and that they were warning potential herbivores of this by a bright autumn display (Hagen et al 2004).

Round about this time (2002), I was approached by a young Swiss researcher, Marco Archetti, who knew that I had a plot of sixty bird cherry trees that I had planted up when I arrived at Silwood in 1992, originally designed to follow-up some work that I had begun whilst at the Forestry Commission looking at the effects of early season defoliation on subsequent tree growth (Leather, 1993, 1995).  Marco convinced me that I had the ideal set-up to test the ‘signalling hypothesis’ and what was to be a very fruitful collaboration began.

We counted arriving gynoparae and their offspring (oviparae) throughout October (Marco making trips over from Oxford where he was then based**) noting leaf colour before and after each count.  As with my PhD work we found that the greener trees were preferentially colonised by the gynoparae and that more oviparae were produced on those trees and that given what I had found earlier that bird cherry aphid gynoparae chose trees that are good hosts in spring (Leather, 1986), Marco felt that we were able to support the honest signalling hypothesis (Archetti & Leather, 2005).  I was slightly less comfortable about this, as there are only two species of aphid that attack bird cherry and one of those is very rare and the original signalling hypothesis was based on the premise that it was trees that were attacked by a lot of aphid species that used the red colouration as a keep clear signal.  Anyway, it was published 🙂

That said, others agreed with us, for example, Schaefer & Rolshausen (2006) who called it the defence indication hypothesis, arguing that bright colours advertise high levels of plant defence and that the herbivores would do well to stay away from those plants displaying them. On the other hand, Sinkkonen (2006) suggested that reproductively active plants produce autumn colours early to deter insects from feeding on them and thus reduce their seed set.

Chittka & Döring (2007) on the other hand, suggested that there is no need to look further than yellow carotenoids acting as integral components of photosynthesis and protection against light damage and red anthocyanins preventing photo-inhibition (Hoch et al., 2001) as to why trees turn colourful in autumn.  In other words, nothing to do with the insects at all.  A couple of years later however, Thomas Döring and Marco got together with another former colleague of mine from Silwood Park, Jim Hardie, and changed their minds slightly.  This time, whilst conceding that red leaves are not attractive to aphids but noting that yellow leaves are even more attractive than green ones, suggested that the red colour could be being used to mask yellow (Döring et al., 2009).

Others have their own pet theories.  In recent years, veteran Australian entomologist Tom White has become interested in the concept of insect species that specifically feed on senescent plant tissue (White, 2002, 2015) and added to the debate by suggesting that aphids in general are senescence feeders and thus choose green and yellow as they have longest time to live and that the red leaves are also nitrogen depleted (White, 2009) which is supported by my PhD data (Figure 1).  This resulted in a spirited response by Lev-Yadun & Holopainen (2011) who claimed that he had misunderstood the scenario in thinking that leaves go sequentially from green to yellow to red, which they suggest is rare (I question this) and that actually in trees that go from green to red, the leaves still contain significant amounts of nitrogen, so a deterrent signal is still required.

Maple, green to yellow in this case

Spindle, Euonymus europaeus, green to red

What about those trees and other plants that have red or purple leaves in the spring or all year round and not just in autumn?

Some trees have red foliage all year

Trees like some of the ornamental cherries or copper beech? I haven’t been able to find any papers that suggest that red or purple-leaved varieties of beech and cherries are less susceptible to aphid attack.  My own observations, probably imperfectly recalled, are that copper beech is regularly infested by the beech woolly aphid, Phyllaphis fagi , and just as heavily, if not more so than the normal green-leaved  beech trees.  That of course may just be a reflection that the white waxy wool covering the aphid stands out more against the red leaves.  Perhaps someone out here might like to check this out?  Some work that my friend and former colleague, Allan Watt, (sadly unpublished) did many years ago in Scotland looking at the effect of beech species and cultivar on infestation levels by the beech leaf mining weevil, Rhynchaenus fagi, did not indicate any differences between copper and green cultivars.  It does seem however, that in cabbages, leaf colour can tell the specialist cabbage aphid, Brevicoryne brassciae, if plants are well defended or not, the bluer the cabbage, the nastier it is (Green et al, 2015).

To summarise:

1. Red leaves are produced by the trees in autumn to reduce ultraviolet damage and protect metabolic processes in the leaf.
2. Red leaves are deliberately produced by the tree to warn aphids that their leaves are well defended – honest signalling.
3. Red leaves are produced by the tree to ‘fool’ the herbivores that the leaves are likely to drop soon and warn them to keep away so as to safeguard their fruit – dishonest signalling.
4. The tree is blissfully unaware of the aphids and the aphids are exploiting the intensity of the autumn colours produced by the trees to select which are the best trees to colonise in terms of nutrition and length of time left on the tree.

As I write, the debate still goes on and we seem no nearer to arriving at a definitive answer to the riddle of why trees produce bright leaves in autumn.  If nothing else however, the debate has generated a lot of interest and enabled people to sneak some amusing titles into the scientific literature.  Do make the effort to read the titles of some of the references below.

References

Archetti, M. (2009) Phylogenetic analysis reveals a scattered distribution of autumn colours. Annals of Botany, 103, 703-713.

Archetti, M. & Leather, S.R. (2005) A test of the coevolution theory of autumn colours: colour preference of Rhopalosiphum padi on Prunus padus.  Oikos, 110, 339-343.

Chittka, L. & Döring, T.F. (2007) Are autumn foliage colors red signals to aphids? PLoS Biology , 5(8): e187. Doi:10.1371/journal.pbio.0050187.

Dixon, A.F.G. (1963) Reproductive activity of the sycamore aphid, Drepanosiphum platanoides (Schr) (Hemiptera, Aphididae). Journal of Animal Ecology, 32, 33-48.

Döring, T.F., Archetti, M. & Hardie, J. (2009) Autumn leaves seen through herbivore eyes.  Proceedings of the Royal Society London B., 276, 121-127.

Furuta, K. (1986) Host preferences and population dynamics in an autumnal population of the maple aphid, Periphyllus californiensis Shinji (Homoptera: Aphididae). Zeitschrift fur Angewandte Entomologie, 102, 93-100.

Green, J.P., Foster, R., Wilkins, L., Osorio, D. & Hartley, S.E. (2015) Leaf colour as a signal of chemical defence to insect herbivores in wild cabbage (Brassica oleracea).  PLoS ONE, 10(9): e0136884.doi:10.1371/journal.pone.0136884.

Hagen, S.B. (2004) Autumn coloration as a signal of tree condition. Proceedings of the Royal Society London B, 271, S184-S185.

Hamilton, W.D. & Brown, S.P. (2001) Autumn tree colours as handicap signal. Proceedings of the Royal Society London B, 268, 1489-1493.

Hoch , W.A.,  Zeldin, E.L. & McCown, B.H. (2001) Physiological significance of anthocyanins during autumnal leaf senescence. Tree Physiology, 21, 1-8.

Holopainen, J.K. & Peltonen, P. (2002) Bright colours of deciduous trees attract aphids: nutrient retranslocation hypothesis.  Oikos, 99, 184-188.

Leather, S.R. (1981) Reproduction and survival: a field study of the gynoparae of the bird cherry-oat aphid, Rhopalosiphum padi (L.). Annales Entomologici Fennici, 47, 131-135.

Leather, S.R. (1982) Do gynoparae and males need to feed? An attempt to allocate resources in the bird cherry-oat aphid Rhopalosiphum padi.  Entomologia experimentalis et applicata, 31, 386-390.

Leather, S.R. (1986) Host monitoring by aphid migrants: do gynoparae maximise offspring fitness? Oecologia, 68, 367-369.

Leather, S.R. (1993) Early season defoliation of bird cherry influences autumn colonization by the bird cherry aphid, Rhopalosiphum padi. Oikos, 66, 43-47.

Leather, S.R. (1995) Medium term effects of early season defoliation on the colonisation of bird cherry (Prunus padus L.). European Journal of Entomology, 92, 623-631.

Leather, S.R. & Dixon, A.F.G. (1981) Growth, survival and reproduction of the bird-cherry aphid, Rhopalosiphum padi, on its primary host. Annals of Applied Biology, 99, 115-118.

Leather, S.R. & Lehti, J.P. (1982) Field studies on the factors affecting the population dynamics of the bird cherry-oat aphid, Rhopalosiphum padi (L.) in Finland. Annales Agriculturae Fenniae, 21, 20-31.

Lee, D.W. & Gould, K.S. (2002) Anthocyanins in leaves and other vegetative organs: An introduction. Advances in Botanical Research, 37, 1-16.

Lev-Yadun, S. & Holopainen, J.K. (2011) How red is the red autumn leaf herring and did it lose its red color? Plant Signalling & Behavior, 6, 1879-1880.

Merzlyak, W.N. & Gittelson, A. (1995) Why and what for the leaves are yellow in autumn? On the interpretation of optical spectra of senescing leaves (Acer platanoides L.). Journal of Plant Physiology, 145, 315-320.

Sanger, J.E. (1971) Quantitative investigations of leaf pigments from their Inception in buds through autumn coloration to decomposition in falling leaves.  Ecology, 52, 1075-1089.

Schaefer, H.M. & Rolshausen, G. (2006) Plants on red alert – do insects pay attention?  BioEssays, 28, 65-71.

Sinkkonen, A. (2006) Do autumn leaf colours serve as reproductive insurance against sucking herbivores?  Oikos, 113, 557-562.

Stiles, E.W. (1982) Fruit flags: two hypotheses. American Naturalist, 120, 500-509.

Ward, S.A., Leather, S.R., & Dixon, A.F.G. (1984) Temperature prediction and the timing of sex in aphids. Oecologia, 62, 230-233.

White, T.C.R. (2003) Nutrient translocation hypothesis: a subsect of the flush-feeding/senescence-feeding hypothesis. Oikos, 103, 217.

White, T.C.R. (2009) Catching a red herring: autumn colours and aphids. Oikos, 118, 1610-1612.

White, T.C.R. (2015) Senescence-feesders: a new trophic subguild of insect herbivore. Journal of Applied Entomology, 139, 11-22.

Wilkinson, D.M., Sherratt, T.N., Phillip, D.M., Wratten, S.D., Dixon, A.F.G. & Young, A.J. (2002) The adaptive significance of autumn colours.  Oikos, 99, 402-407.

 

 *for a detailed account of the wonderful terminology associated with aphid life cycles read here

**coincidentally he is now a Lecturer at the University of East Anglia in the same Department where I did my PhD

The Seven Ages of an Entomologist – Happy 60th Birthday to Me

Today I turned 60 – an event which has come as a bit of a surprise to me as inside I still feel about 17 😉 I thought, given the occasion and the fine example set by Jeff Ollerton‘s recent birthday blog post  that it seems a good time to reflect on my career in particular and academic careers in general. Despite there already being at least two other excellent articles about the “Seven Ages”, Jerry Coyne’s, The Seven Ages of the Scientist and Athene Donald’s The Seven Ages of an Academic Scientist, I felt no qualms in adding my own modest contribution to the genre 😉

Given my own career trajectory it turns out that I need more than seven ages, so as an entomologist I feel justified in adding five larval or nymphal instars to the traditional progression.

 

The Larval Stages

The Infant (first instar)

According to Shakespeare “mewling and puking in the nurse’s arms”, which spending the early part of my childhood in colonial Ghana is actually very apt,

although the photograph below shows a very contented baby indeed.

I have no entomological memories from this time, although given that then it was normal practice to leave babies outside in their prams, I am sure that I was exposed to the whole range of flying Ghanaian insects. There is some evidence of an early interest in nature and entomology in the picture below where I seem to be investigating a small white butterfly whilst indulging in some early forestry work.

My first real biological memory, is however, non-entomological, the blue whale skeleton in the Natural History Museum London in 1958 when my parents were on home leave.

 

The Schoolboy (second instar)

 In 1960 my father was moved to Jamaica to work in the Department of Agriculture as a Plant Pathologist and this is where I started my formal education. Shakespeare describes the schoolboy as “whining schoolboy with his satchel, and shining morning face, creeping like a snail unwillingly to school”.

I certainly had a satchel and it is from this period of my life that I have my first definite entomological memories. We lived in a suburb of Kingston, 32 Gardenia Avenue in Mona Heights. My father kept bees and I spent a lot of time playing with ants, conducting behavioural experiments with crab spiders and having close encounters with wasps and apparently in this picture from 1961, helping with my father’s very luxuriant garden; he grew a great variety of ornamental plants as well as fruit and

     

 

vegetables, including grapes, bananas, passion fruit, papayas, peanuts and breadfruit as well as coffee and more traditional vegetables. My final school report from my time in Jamaica shows a prescient comment from my biology teacher;

 

Secondary school (third instar)

My father’s next posting was to Hong Kong to work for the Ministry of Agriculture; his office was in the New Territories but we lived in Kowloon (Wylie Gardens) where I attended King George V School. Biology was again my favourite subject but apart from cockroaches and ants my entomological experiences were very limited.

           Before braces – 1966

Keeping my mouth shut to hide my orthodontic appliances 1968.

 

Boarding school (fourth instar)

In 1968 my father returned briefly to the UK before his next posting to Fiji and I was sent to a state school, Ripon Grammar School, which had a boarding section. I was to spend five relatively happy years there and despite the competing interests of girls and sports, further developed my interest in invertebrate zoology, due in the main part to my zoology teacher ‘Brian’ Ford. I have many happy memories of pond dipping, searching for Cepea nemoralis and generally fossicking around in hedgerows.

When on school holidays in Fiji I found time to investigate the local insect and amphibian fauna; our house seemed to attract toads in huge numbers which my brothers and I used to competitively collect in buckets for later release.

 

Sixth form (final instar)

In my two final years at school sport and girls continued to play a larger part in my life than entomology although I see from the fly-leaf of my books from that time that I owned and had read both volumes of Ralph Bucshbaum’s Life of the Invertebrates and also Darwin’s Origins.

Ripon Grammar School 2^nd XV – I am third from the left on the front row.

 Careers advice when I was at school was not very sophisticated and if you did Biology ‘A’ Level and were a school prefect, it was automatically taken that you were either destined to be a Doctor, a Vet or a Dentist.

I was no different and despite my misgivings, duly applied for and was accepted at Birmingham University to read Medicine. As luck would have it, things did not work out as planned and after a less than happy year at Aston University in Birmingham, in 1974 I left Birmingham and moulted into a proto-entomologist at the University of Leeds.

 

The Undergraduate

The discovery that learning can be fun and that there might actually be a career in doing something that you enjoy.

I did a now extinct degree (although I have plans to exhume it), Agricultural Zoology, essentially a year of vertebrate zoology, with two years of invertebrate zoology, essentially applied entomology, parasitology and nematology. I loved it and thrived on it and grew my hair even longer.

I decided to become an entomologist in my second year and discovered the wonders of aphids at the same time. It was also round about this time that I decided I was going to become a university academic and started to work a lot harder; the logical end point of someone with a mother who was a secondary school biology teacher and a father who was a research scientist.

 

The Postgraduate

Discovering that being on “the road to find out” (Cat Stevens) is exhilarating

I did my PhD at the University of East Anglia in Norwich – Aspects of the Ecology of the Ecology of the Bird Cherry Aphid, under the supervision of Professor Tony Dixon. A totally fantastic time, despite the ‘second year blues’ which all PhD students seem to go through when they think that they don’t have enough data. I was lucky enough to be in a large research group, at one stage there were thirteen of us in the lab, so there was always plenty of help and advice available. In addition we had the excitement of conferences and the first unsteady steps towards learning to lecture, mainly demonstrating in undergraduate practicals; I spent a lot of time pithing frogs for physiology classes (don’t ask) and also tutoring first year students in mathematics. We also played a lot of squash and enjoyed our social life; for those of you who know Norwich, The Mitre pub on Earlham Road, was our regular haunt.

 

The post-doc

Discovering how to run a research lab

I did two brief post-docs, the first in Finland, under the auspices of the Royal Society and the

second back at the University of East Anglia funded by the Agriculture and Food Research Council, both working on cereal aphids. At this stage of my career I started to learn how to supervise postgraduate students; the first port of call in a busy lab after the senior PhD student has failed to supply an answer is always the post-doc as the lab head is inevitably very busy. I also got my first real opportunity to lecture undergraduates, which turned out to be a lot harder than I had thought it would be even when talking about my own research.

 

Interlude or host alternation

 The Research Scientist

 Discovering that directed research on its own is not enough

In a normal academic career, the next stage after post-doc is an appointment as a University Lecturer. In the early 1980s university lectureships were in short supply and many of us who would normally have gone into an academic career found ourselves either having to go abroad as lecturers at Commonwealth universities (I was offered but turned down a lectureship at Kano University in Nigeria) or joining research institutes. In 1982 I joined the UK Forestry Commission’s Northern Research Station where I spent ten years as a forest entomologist, answering enquiries, conducting directed research and giving the occasional guest lecture. I was however, lucky enough to be able to gain some PhD supervisory experience and after ten years, the last five which were increasingly frustrating, was lucky enough in 1992 to be appointed to a Lectureship at the Silwood Park campus of Imperial College.  In retrospect this was the last time I was able to spend about 90% of my time at the bench and in the field doing ‘hands on’ research, but I have never regretted moving into academia – the opportunity of being able to pass on what you have discovered and hopefully enthuse and motivate a new generation more than makes up for the loss.
Back to the primary host

 

The Lecturer

When I discover that I love teaching

You may have noticed that I have had a haircut; it was a source of some amusement to me that on joining the university sector I was expected to get my hair cut.

I was appointed as a Lecturer in Pest Management to teach on the world-renowned MSc Entomology course at Silwood Park, and as I was replacing a specific person (Geoff Norton), although not in exactly the same subject area, my ‘grace’ period was shorter that it might have been. Normally at research intensive institutions like Imperial College, new appointments are given two to three years to apply for grants and get their research groups started before being given teaching and departmental jobs. I had a year, but as I discovered that I very much enjoyed teaching (something that many of my colleagues then and later found very strange) I was not dismayed. Unlike some of my colleagues I had read the dictionary definition of the word lecturer: noun. One who delivers lectures, especially professionally.   I have never really understood the mentality of those who aspire to university positions and yet find the idea of having to teach students not only a distraction but in some cases abhorrent and to be avoided at all costs and strive to obtain funding to buy them out of teaching as soon as possible. Some of my senior colleagues at Imperial College (and elsewhere) had and have almost no experience of teaching at all and so have no idea of what is involved in delivering a decent course, a state of affairs that explains some of the very strange decisions that are made at some of the research intensive universities in the UK.   I often felt that they would be much happier in a research institute.

I also discovered that if you take teaching seriously then your ‘bench time’ is much reduced and you begin your career as a research manager, appointing PhD students and post-docs to carry your research ideas forward. I made a decision early on that I would attempt to keep some of my skills extant and set up a long-term field project looking at the insect communities living on sycamores at Silwood Park, especially the aphids. This meant that I had to set a day a week aside to collect data. By doing this it meant that I had a reality check on what was actually possible. I have seen too many colleagues who because of the time they had spent away from the bench or the field, had totally unrealistic expectations of what was actually possible to be achieved by their students and research assistants.

 

The Senior Lecturer

When the Department discovers that I love teaching

In 1996 I was promoted to Senior Lecturer (I think that it is a real shame that some UK universities have decided to adopt North American terminology and introduce the title of Associate Professor, apparently to avoid confusing the rest of the World. At Imperial College promotion to Senior Lecturer was to reward teaching excellence and was usually the kiss of death for any further promotion.

Senior Lecturer in Applied Ecology

 I was as well as teaching on the MSc Entomology course doing an increasing amount of undergraduate teaching including a final year course in Applied Ecology of which I was very proud, hence the decision to retitle myself. I was also very busy with external activities, being on the Editorial Board of the Bulletin of Entomological Research and just been appointed as Editor-in-Chief of Ecological Entomology, just finished a term on the council of the Royal Entomological Society and been appointed to a slew of Departmental and University committees. My research group was really starting to take off, I was supervising 8 PhD students at the time; given the poor return rate on major grant applications in the UK, I decided early on that going for PhDs was a better use of my limited time and this is a strategy that I have mainly followed to the present day.

This does not include MSc or BSc students – they would add about 10 to each yearly figure from 1995 onwards

The Reader

 When I discover that it is possible to get even busier

In 2002 I was promoted to Reader one of the definitions of which according to Chambers’s Twentieth Century Dictionary is defined as follows; Old English rǣdere ‘interpreter of dreams, reader’. In the UK university system, it is the rank below full Professor and comes with an endowed title, in my case I chose to become Reader in Applied Ecology to reflect the

myriad teaching roles I had accumulated and also to encompass the fact that my research group no longer dealt solely with arthropods, vertebrates had somehow sneaked their way in. Looking at Athene Donald’s list I see that I was pretty much doing a professorial role, serving on external committees, validating degrees for other universities and acting as an external examiner. I was also appointed as Editor-in-Chief of Insect Conservation and Diversity, a new journal for the Royal Entomological Society. My administrative duties had also continued to increase.  It was no wonder that my beard was getting greyer! I was however still preparing my own talks, although I will confess that a lot of my data analysis was being passed on to members of the group, duly acknowledged of course. I am extremely grateful that I have always had a loyal and very supportive research group, without their help life would have been impossible.  My thanks to you all (if any of you are reading this).

 

The Professor

Discovering the joys of being pretty much able to do what I want (with certain restrictions)

It became increasingly obvious that things could not carry on as they were, my teaching and administrative loads were becoming ridiculous; our Director of Teaching calculated that I was actually doing more teaching than anyone else in the Department including the Teaching Fellows. I was seriously considering early retirement although I was reluctant to do this as I was sure that with my retirement the last entomology degree in the UK would quickly disappear. Luckily in 2012 my team and I were miraculously offered the chance to move to a new more supportive location, Harper Adams University in Shropshire.

So now I have become a Senior Professor, with a new entomology building, with less undergraduate teaching, which I miss, and a role that requires me to sit on more external and internal committees, to meet the great and the good and to make solemn pronouncements.  At the same time however, it does allow me to plough my own furrow and to influence university policy. Most importantly I no longer feel that I am beating my head against a brick wall and that the future of entomology as a degree course in the UK is much safer than it was five years ago.  I think I am at Stage 4 in Jerry Coyne’s list as I now find that I am much more interested in synthesizing and disseminating what I have learnt rather than doing original research – I can feel a book coming on 😉

My hope is that in five years time when I become a retired Professor and my hair and beard colour are the same, that entomology will be taught at more than one university in the UK and not just at postgraduate level.

A small point of personal satisfaction, is that, despite my elevation, I still do not own a suit 😉

 

For reference

Jerry A. Coyne’s summary, reproduced from his blog

1. As student, listens to advisor give talk on student’s own work
2. As postdoc, gives talks about his/her own work
3. As professor, gives talks about his/her students’ work
4. Talks and writes about “the state of the field”
5. Talks and writes about “the state of the field” eccentrically and incorrectly—always in a self-aggrandizing way.
6. Gives after-dinner speeches and writes about society and the history of the field
7. Writes articles about science and religion

 

And the famous original from which the title is borrowed and adapted.

 

Seven Ages Of Man

(from As You Like It by William Shakespeare)

All the world’s a stage,

And all the men and women merely players,

They have their exits and entrances,

And one man in his time plays many parts,

His acts being seven ages. At first the infant,

Mewling and puking in the nurse’s arms.

Then, the whining schoolboy with his satchel

And shining morning face, creeping like snail

Unwillingly to school. And then the lover,

Sighing like furnace, with a woeful ballad

Made to his mistress’ eyebrow. Then a soldier,

Full of strange oaths, and bearded like the pard,

Jealous in honour, sudden, and quick in quarrel,

Seeking the bubble reputation

Even in the cannon’s mouth. And then the justice

In fair round belly, with good capon lin’d,

With eyes severe, and beard of formal cut,

Full of wise saws, and modern instances,

And so he plays his part. The sixth age shifts

Into the lean and slipper’d pantaloon,

With spectacles on nose, and pouch on side,

His youthful hose well sav’d, a world too wide,

For his shrunk shank, and his big manly voice,

Turning again towards childish treble, pipes

And whistles in his sound. Last scene of all,

That ends this strange eventful history,

Is second childishness and mere oblivion,

Sans teeth, sans eyes, sans taste, sans everything.

The Dead Entomologists Society

The late and the great: who are the most influential dead entomologists?

In the run-up to the Christmas holidays an ex-student of mine, Andy Salisbury, now at RHS Wisley, and I were discussing who were the most influential dead entomologists ever.  We had begun our discussion discussing Harold Maxwell-LeFroy, http://en.wikipedia.org/wiki/Harold_Maxwell-Lefroy, who among other claims to fame, was the first editor of Annals of Applied Biology http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1744-7348 , the founder of Rentokil and former Professor of Entomology at Imperial College.  He was also famous for having killed himself accidentally whilst trying out a pesticide.

Up until the early part of this century he was commemorated in the Biology Department at Imperial College’s Silwood Park Campus with a laboratory named after him; sadly with the move into the new Hamilton Building this no longer exists.

We agreed that the great Alfred Russel Wallace was the most influential entomologist of all time; we felt that claiming Darwin as an entomologist, although famous for his beetle collecting, and despite being a Fellow of the Royal Entomological Society, might be a step too far.  So after Wallace, who was the most influential dead entomologist?

I think that this depends on how you define influential – for entomologists of my generation and the one before us, i.e. those born between 1930 and 1960; Imms and Wigglesworth are probably the two who most influenced us mainly because they were our recommended undergraduate texts  (those were the days when you could do entomology  in the UK as an undergraduate);

today I guess these have been replaced by Chapman’s The Insects: Structure and Function and Gullan & Cranston’s Outline of Entomology.  But of course outside the field of entomology who remembers Imms and Wigglesworth?  So we should, I think, be looking for entomologists whose influence has extended more widely;  a slightly tongue in cheek contender would be Thomas Moufet,  http://en.wikipedia.org/wiki/Thomas_Muffet who is possibly apocryphally, remembered for the nursery rhyme about his daughter Little Miss Muffett and more substantiated, for his compilation of the Theatre of Insects, but he was mainly a man of

medicine and more interested in spiders than insects in general.  A contender widely known outside the entomological world would be H W Bates, http://en.wikipedia.org/wiki/Henry_Walter_Bates#Taxonomy who is today remembered in the term Batesian

mimicry  http://en.wikipedia.org/wiki/Mimicry#Batesian ; as a former President of the Royal Entomological Society (1868-1869) I think that we can safely claim him as an entomologist.

Other entomologists that have had an influence  on the rest of the non-entomological world, albeit not widely known outside the biological or medical sciences are Fabre and his Lives of  various insects http://en.wikipedia.org/wiki/Jean-Henri_Fabre , Karl von Frisch http://en.wikipedia.org/wiki/Karl_von_Frisch for his work on honeybees, especially in deciphering the waggle-dance,  E B Ford for his work on ecological genetics and for inspiring Kettlewell’s work on Biston betularia,  http://en.wikipedia.org/wiki/E._B._Ford  and more recently Richard Southwood  http://en.wikipedia.org/wiki/Richard_Southwood   (Methods in Ecology) but initially a Hemipteran specialist and Miriam Rothschild  http://en.wikipedia.org/wiki/Miriam_Rothschild for her work on a range of entomological subjects but very famously for her work on fleas.

The acid test of course, is how many living entomologists the man or woman in the pub can bring to mind when asked, let alone those that have joined the Dead Entomologist’s Society. Perhaps it is the fate of entomologists to be largely overlooked, much like the small but highly important organisms we work on 😉

I would dearly love to hear your thoughts on which the most influential dead entomologists are.  I am well aware, that my list is very Euro- and UK-centric.  So let’s see some nominations from the rest of the world, suitably justified of course!

Post script

Getting my vote as a contender for an American entomologist of great influence, albeit British-born, would be Charles Riley, sometimes known as the Father of Biological Control http://en.wikipedia.org/wiki/Charles_Valentine_Riley , but then again, who outside the world of entomology has heard of him?