Amateur, professional and academic interactions: A day with the British Entomological and Natural History Society

Towards the end of last year I was asked by Claudia Watts, the President of the British Entomological and Natural History Society http://www.benhs.org.uk/site/?q=node/1 if I would like to be nominated to become a member of their Council. I thought about it for a while and then said yes – my resolution to learn to say NO has sadly not been very successful 😉

There was one snag though, I was not a member of the BENHS; that was, however, easily remedied by completing an application form and being duly approved by the membership secretary and then paying my annual subscription of £19. This process made me think, why wasn’t I already a member of the BENHS? The aims of the Society “are the promotion and advancement of research in entomology with an increasing emphasis now being placed on the conservation of the fauna and flora of the United Kingdom and the protection of wildlife throughout the world”, which is something I have been interested and involved with for almost forty years.

Until I graduated from Leeds the only societies I had belonged to were university ones. On starting my PhD I joined the Association of Applied Biologists, the British Ecological Society, the then Institute of Biology (my father was a founder member) and the Royal Entomological Society. All these had one thing in common, they were, for want of a better description, professional graduate societies, although the Royal Entomological Society did, and does have non-academic and non-professional Members and Fellows. So I guess there was a bit of snobbery involved – the British Entomological and Natural History Society despite its distinguished past, founded 1872, and publishing a quarterly journal and a variety of important identification guides, was in my world view at the time, akin to the Amateur Entomological Society, founded in 1935. A reading of the aims of the AES would however, have told me otherwise, “Our objective is to promote the study of entomology, especially amongst amateurs and the younger generation.”

The objectives of the BENHS, originally the South London Entomological and Natural History Society are remarkably similar to those of the Royal Entomological Society (originally of London), the main differences now, being their financial resources and scale of activities. So having entered an academic career and eventually ending up in one of the UK’s top research intensive universities I had no incentive or inclination to join a society or organisation that did not instantly suggest a professional affiliation, and in the case of the Institute of Biology (now Society of Biology) and the Royal Entomological Society giving me additional letters after my name!

That said, a significant proportion of the UK academic entomological community have published and continue to  publish in the less ‘high impact’ end of the journal spectrum (e.g. Southwood & Johnson, 1957; Dixon, 1958) and indeed me too (Leather & Brotherton, 1987; Leather, 1989). This has been more fully documented elsewhere (Hopkins & Freckleton, 2002). The need for such journals is perhaps not fully appreciated. One of my former students, a Hemipterist of some note, who has published extensively in this journal stratum used to leave these off his job applications until he was asked at a post-doc interview if he had published in these journals as they considered it an important requisite of the job in question as it involved a lot of field work and insect identification. Without journals such as these published here and elsewhere, much of the basic knowledge needed for academic entomologists, ecologists, zoologists etc. to conduct their research would be greatly hampered.

So how does it work in practice? I duly turned up for my first BENHS AGM on Saturday 21^st March at the very apposite venue, the Oxford University Museum of Natural History, in time

for the pre-meeting mingle with coffee and biscuits and met some of my former MSc students, now doing PhDs, old friends, current MSc students,  undergraduates, one of whom I had interviewed the day before for a place on the MSc in Entomology that I run, and even a scattering of much younger entomologists.  So a real mixture of amateurs, professionals (those making a living from entomology but not employed by a research institute or university) and current and retired academics.

The programme included a mixture of entomologists, ranging from Matt Shardlow, CEO and founder of Buglife, entomological consultant and saproxylic beetle guru, Keith Alexander, PhD student Tom Wood from Sussex University talking about his pollinator research which has just made the news, author and professional entomologist Richard Jones plugging his latest book and school boy Louis Guillot talking about his research into ant colony development; many of which he stores beneath his bed. We also had an entertaining Presidential Address by Claudia Watts about insects and art through the ages.

Having the AGM in the museum was also of course a very positive plus as in the breaks I was able to have a quick look at some of the exhibits – this one particularly caught my eye, although my camera and photographic skills do not do it justice.

 

My take home message is that a) I was very foolish not to have got involved with the BENH many years ago and b) given the problems we have in getting the media and general public to understand the importance of entomology and insects to the well-being of the world, that many more of us pre-retirement academic entomologists should be willing to get involved with this and other similar societies.

References

Dixon, A.F.G. (1958) The protective function of the siphunculi of the nettle aphid, Microlophium evansi (Theob.). Entomologist’s Monthly Magazine, 94, 8.

Hopkins, G.W. & Freckleton, R.P. (2002) Declines in the numbers of amateur and professional taxonomists: implications for conservation. Animal Conservation, 5, 245-249.

Leather, S.R. (1989) Phytodecta pallida (L.) (Col.,Chrysomelidae) – a new insect record for bird cherry (Prunus padus). Entomologist’s Monthly Magazine, 125, 17-18.

Leather, S.R. & Brotherton, C.M. (1987) Defensive responses of the pine beauty moth, Panolis flammea (D&S) (Lepidoptera: Noctuidae). Entomologist’s Gazette, 38, 19-24.

Southwood, T.R.E. & Johnson, C.G. (1957) Some records of insect flight activity in May, 1954, with particular reference to the massed flights of Coleoptera and Heteroptera from concealed habitats. Entomologist’s Monthly Magazine, 93, 121–126

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 Verrall Supper 2015 – A Photographic Record

Wednesday March 4^th 2015 was the occasion of the latest Verrall Supper, an annual event hosted by the Entomological Club, the oldest extant entomological society in the world.  I am, for my sins, the Supper Member, which means that I have to organise the event, for a more detailed description of my role click here.  This year we continued our association with The Rembrandt Hotel as they had done such a good job last year and the year before.  This year almost all the invitations were sent by email and despite the 14% increase in suggested subscription to £48, we had a very good response; as I pointed out to one of the students, show me somewhere else in central London where you can get a three course dinner with coffee and  half a bottle of wine, plus the company of so many entomologists!

In the end we had 181 guests, 55 of whom were female, last year we only had 46 female members so we are definitely moving in the right direction, although I am still keen to get equal numbers.  My impression was that the average age of the membership is definitely decreasing which can only be a good thing.  Enough writing I think, let the photographs speak for themselves.

Jim Hardie with Clive Farrell of the Entomological Club – once again my thanks to Clive for helping man the Reception Desk

A mixed bag of entomologists enjoying good conversation whilst waiting for the main course

All ex- or present students of mine, Katy Reed, Lauren Fuller, Jen Banfield-Zanin, Mark Ramsden, Aislinn Pearson

Andrew Salisbury, RHS Wisley, holds forth

 

Ashleigh Whiffin and Craig Perl recreate last year’s photo

Can you spot the Coleopterist?

Entomologists with beer

Garth Foster making a point

Gemma Hough and James Hourston

Did you know that Hagrid was an entomologist?  Actually Richard Comont

Can you spot Helen Roy, the newest member of the Entomological Club?

Three ex-MSc students – Jade Taylor, Linda Birkin and Laurence Livermore and Hillery Warner.

Mainly current Harper Adams MSc students – Josh Jenkins Shaw, Chris Mackin, Andy Cutts, Aidan Thomas, Richard Prew, Kelleigh Greene, Jordan Ryder (now a PhD student) Dave Stanford-Beale

Marion Gratwick has attended more Verrall Suppers than any other female member.

Mark Ramsden, Adrian De Palma, Jen Banfield-Zanin, Gemma Hough

More young entomologists, Ailsa McLean, Paul Manning, Chris Jeffs and James Hourston

John Badmin centre stage

Some older entomologists

Linda Birkin, Laurence Livermore, Hillery Warner, Aurora Sampson

Some hymenopterists, including Mark Shaw and Charles Godfray

Some Royal Entomological Society worthies including Gordon Port, Jim Hardie and Archie Murchie

On the top table, Stephen Clement who travelled all the way from the USA, speaking to Gill van Emden

The top table, Gill van Emden, Stephen Clement, Van, Clive Farrell, Chris Lyal, another overseas visitor Junhao Huang, my jacket, Camille Parmesan, Richard Harrington and Tilly Collins out of shot.

Has Richard Harrington made Tilly Collins cry?  Are his jokes really that bad?

The Verrall lecturer, Sue Hartley in deep conversation with Hugh Loxdale and Peter Leckstein.  Unfortunately she couldn’t stay for the Supper.

Not all entomologists are male and bearded, Ruth Carter, Kirsten Miller, Fran Sconce, Will Nash, Nathan Medd , Hannah Wickenden, Jasper Hubert

More young entomologists Joe Nunez, Ricahrd Comont, Amo Spooner, Katy Dainton, Molly Carter, Sally-Ann Spence

Some of the older entomologists

and finally

The Supper Organiser looking a little bit stressed!

I look forward to seeing all of you next year on Wednesday 2^nd March.

Ten Papers that Shook My World – Haukioja & Niemelä (1976) – the plant “immune response”

To me this is a landmark paper, both personally and for ecology in general.   I first came across it in the second year of my PhD at the University of East Anglia (1978) and given where it was published, would probably never have seen it if my supervisor, Tony Dixon, hadn’t had a collaborative link with Erkki Haukioja of Turku University (Finland).

That individual plants of the same species are more or less susceptible (constitutive or innate resistance) to pests and diseases has been known for a very long time (e.g. Painter, 1958; Beck, 1965) and has been exploited by plant breeders as part of many pest management programmes.  Despite the stunning footage of the questing bramble in David Attenborough’s classic documentary The Private Life of Plants, plants are often thought of as passive organisms.  The idea that plants might actually respond directly and quickly to insect attack was more in the realms of science fiction than science fact, but this all changed in the 1970s. In 1972 a short paper in Science (Green & Ryan, 1972) suggested that plants might not be as passive as previously thought. Green & Ryan working in the laboratory with the Colorado Potato Beetle, Leptinotarsus decemlineata, showed that when tomato leaves were damaged by beetle feeding the levels of a proteinase inhibitor were raised not just in the wounded leaves but in nearby leaves as well. As proteinase inhibitors were well-known to be part of the plant defence system, they hypothesised that this was a direct response of the plant to repel attack by pests and that it might be a useful tool in developing new pest management approaches. So what does this have to do with two Finnish entomologists?

Erkki Haukioja and his long-term collaborator, Pekka Niemelä were working on an important lepidopteran defoliator of birch, in the far north of Finland, at the Kevo Subarctic Research Station.

http://www.eu-interact.org/field-sites/finland-4/kevo/

The defoliator that they were working on was the autumnal moth, now Epirrita autumnata, but then Oporinia autumnata.

http://ukmoths.org.uk/show.php?bf=1797

The autumnal moth, as with many tree-feeding Lepidoptera, has a 7-10 year population cycle (Ruohmäki et al., 2000).

Natural enemies are often cited as the causes of these cycles (Turchin et al., 1999) although other factors such as weather (Myers, 1998) or even sunspot activity (Ruohmäki et al., 2000)

have also been suggested. It had also been suggested that the marked population cycles of the larch bud moth, Zeiraphere diniana were caused by changes in the susceptibility of their host trees after defoliation (Benz, 1974). In 1975, Haukioja and his colleague Hakala, attempting to explain the cyclical nature of the E. autumnata population cycles wondered if they were being driven by the insects themselves causing changes in the levels of chemical defence in the trees. To test this Erkki and Pekka did two neat field experiments, remember Green & Ryan’s work was laboratory based and did not test the effects seen on the insects. They first fed Epirrita larvae on foliage from previously defoliated and undefoliated birch trees and found that the pupae that developed from those larvae fed on previously defoliated trees were lighter than those that had fed on previously undefoliated trees (Hauikioja & Niemelä, 1976). At the same time they also did an experiment where they damaged leaves but then rather than feeding the larvae on those leaves, fed them on nearby adjacent undamaged leaves and compared them with larvae feeding on leaves from trees where no damage had occurred. Those larvae feeding on undamaged leaves adjacent to damaged leaves grew significantly more slowly than those feeding on leaves that came from totally undamaged trees (Haukioja & Niemelä, 1977). So pretty convincing evidence that the trees were responding directly to insect damage and altering their chemistry to become more resistant, i.e. an induced defence and not a constitutive one.

Their results had a major impact on the field. The great and the good from around the world found it a fascinating subject area and a plethora of papers investigating the effects of insect feeding on induced defences in birch and willow trees soon followed (e.g. Fowler & Lawton, 1984a; Rhoades, 1985; Hartley & Lawton, 1987) and not forgetting the original researchers (e.g. Haukioja & Hahnimäki, 1984). I, with the aid of colleagues, also added my ‘two pennorth’ (I did say the idea shook my world) by extending the concept to conifers (Leather et al., 1987; Trewhella et al., 1997). The terms rapid induced resistance and delayed induced resistance soon entered the language, the first to describe those changes that occurred within minutes of feeding damage and the second, those that did not take effect until the following year (Haukioja & Hahnmäki, 1984; Ruohmäki et al., 1992) Such was the interest generated by the topic that by 1989 there were enough studies for a major review to be published (Karban & Myers, 1989).

Controversy reared its ugly head early on when Doug Rhoades suggested that not only did plants resist insect attack actively but that they could talk to each other and warn their neighbours that the ‘bad guys’ were in the neighbourhood (Rhoades, 1983, 1985). This sparked a brief but lively debate (e.g. Fowler & Lawton, 1984b, 1985). Ironically it is now taken as axiomatic that plants talk to each other using a range of chemical signals (van Hulten et al., 2006; Heil & Ton, 2008) as well as informing the natural enemies of the pests that a suitable food source is available (e.g. Edwards & Wratten, 1983; Amo et al., 2013; Michereff et al., 2013).

A great cartoon from Jurriaan Ton at Sheffield University. https://www.shef.ac.uk/aps/staff-and-students/acadstaff/ton-jurriaan

We now have a greatly increased understanding of the various metabolic pathways that induce these defences against different insect pests (e.g. Smith & Boyko, 2007) and can, by genetically manipulating levels of compounds such as jasmonic and salicyclic acids or even applying them directly to plants affect herbivorous insect communities and their natural enemies thus improving crop protection (e.g. Thaler, 1999; Cao et al., 2014; Mäntyllä, 2014). No wonder this was an idea that shook my world, and yours.

 

Post script

The study of induced plant defences or resistance is now dominated by molecular biologists and current practice is to use the term priming and not induced defence. The increased understanding that this new generation has brought to the field is undeniable but I always feel it is a great shame that they seem to have forgotten those early pioneers in the field.

 

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