Christopher John Lamb CBE. 19 March 1950 - Royal …rsbm.royalsocietypublishing.org/content/roybiogmem/56/189.full... · Christopher John Lamb Cbe 19 march 1950 — 21 august 2009

21 August 2009−−Christopher John Lamb CBE. 19 March 1950

Joanne Chory and Rod Casey

originally published online August 18, 2010, 189-213, published 18 August 2010562010 Biogr. Mems Fell. R. Soc.

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Christopher John Lamb Cbe19 march 1950 — 21 august 2009

Biogr. Mems Fell. R. Soc. 56, 189–213 (2010)

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Christopher John Lamb Cbe

19 march 1950 — 21 august 2009

elected Frs 2008

By Joanne Chory1 and rod Casey2,*

1 Howard Hughes Medical Institute and Plant Biology Laboratory of the Salk Institute for Biological Studies, 10010 North Torrey Pines Road, La Jolla,

CA 93207, USA 2 11 Central Crescent, Hethersett, Norwich NR9 3EP, UK

Chris Lamb was one of the most influential plant biologists of his time. he was director of the plant biology Laboratory at the salk institute and of the John innes Centre, two of the world’s leading plant and microbial research institutions. he was recognized by the institute for scientific information as one of the most highly cited researchers worldwide in plant and animal science. his field of study was plant–pathogen interactions, and he made seminal contributions to the understanding of the molecular basis of plant defence. his efforts led to discoveries such as how cell wall composition influences microbial entry, which plant genes are rapidly induced on infection, and the role of reactive oxygen and nitrogen intermediates in plant defence signalling. his work demonstrated a number of links between defence responses in plants and in animals. he believed that plant and microbial science research could be used to improve the world, and he took his belief to the local communities, to regional and national media and to Westminster. he had a skill for talent spotting, and one of his major legacies is the number of careers that he nurtured and are now flourishing at universities and institutes across the world.

The early days

Chris Lamb was born in York on 19 march 1950 and was named malcolm stewart Dixon. shortly afterwards he was adopted by eileen and John Lamb and was renamed Christopher John Lamb. Chris had a happy childhood, being the centre of attention for his parents and small

doi:10.1098/rsbm.2010.0014 191 this publication is © 2010 the royal society

* rod Casey worked closely with Chris Lamb on the John innes Centre management board until his retirement in 2006.



192 Biographical Memoirs

extended family. as a little boy, he spent his summers in a caravan by the sea at saltburn, from where his father took the train to work in middlesbrough every day. his father was secretary to the Constantine merchant shipping company, and when Chris was a teenager he went on several trips as a cabin boy on the company’s ships, visiting eastern europe, italy and scandinavia.

Chris was always a keen sportsman. although his father had little interest in football, Chris’s maternal grandfather took him, his only grandchild, to see both sunderland and middlesbrough matches, thereby fostering in Chris a lifelong passion for middlesbrough Football Club. Chris attended middlesbrough high school from 1961 to 1968; he played football for the school, and cricket for both the school and middlesbrough schools. Chris was nicknamed ‘Charlie’ by his schoolmates because he played centre-half and they thought he emulated Charlie hurley, the famous sunderland and eire international who was one of the finest players ever in that position.

Up to the age of about 15 years, Chris wanted to become an architect, but then he read an article in Scientific American about the medical research Council Laboratory for molecular biology in Cambridge, at the time of max perutz, John Kendrew, Frederick sanger, Francis Crick, sydney brenner and others. he was so excited by what he read that he decided he wanted to go to Cambridge and be a molecular biologist. Chris went on to apply for a place to read natural sciences at Fitzwilliam College. although his school reports at the time sug-gested that he was taking things easy, Fitzwilliam must have recognized his potential, because they offered him a place for october 1969.

During 1968–69 Chris worked at the iCi laboratories in macclesfield, where he had his first taste of professional biochemical research.

CamBridge and oxford

in Freshers’ Week 1969 at Cambridge, Chris attended a book sale organized by a new hall student who was giving up her science course. it was at the tea party after the sale that Chris met Jane Wright, a new hall student, a meeting that was the beginning of an almost 40-year partnership. the couple married in 1970, at the end of Chris’s first year, and went on to have three children: a daughter, Catherine, born in 1971 while they were undergraduates, and two sons, William, born in 1976, and Donald, born in 1985. Chris obtained first-class honours in natural sciences (biochemistry) in 1972.

after graduating, Chris remained in Cambridge and took a phD under the guidance of philip rubery on biochemical studies of the enzyme phenylalanine ammonia-lyase (paL) (figure 1). paL catalyses the first step in the synthesis of phenylpropanoid natural products, and Chris focused on the mechanism underlying the wound-induced synthesis of paL in potato. he was interested to know whether the increase in paL activity after wounding was the result of de novo synthesis or the activation of existing enzyme, and addressed the ques-tion by using various inhibitors of protein and rna synthesis. he also conducted theoretical work on the behaviour of enzymes, such as paL, that exhibit negative rate cooperativity and thereby have implications for biochemical pathway control through feedback inhibition. he was inordinately proud of the fact that this work was published in the Journal of Theoretical Biology (1)*, to the extent that he was still reminding his colleagues of the fact some 30 years

* numbers in this form refer to the bibliography at the end of the text.



Christopher John Lamb 193

later. his approach to the interpretation of quantitative relationships between paL and phenyl-propanoid accumulation (23) was an early example of what would now be known as plant systems biology.

in 1975 Chris joined the University of oxford botany school as an iCi postdoctoral Fellow at st edmund hall (whose football team he captained), working in Vernon butt’s laboratory. by then it was clear that the tools of molecular biology were an essential part of modern biochemistry, and Chris made it his mission to apply them to the aspects of plant biochemistry that interested him. he teamed up with rick Dixon, who was completing his phD working on enzyme induction in response to perceived microbial attack, using elicitor-treated bean cell suspension cultures. the aim was to find whether or not plant cell responses, induced in response to pathogen signalling, were the result of the activation of specific sets of genes. at that time, it was no mean task to distinguish between the activation of pre-existing defences and the de novo synthesis of rnas, proteins and antimicrobial compounds: there

Figure 1. Chris in the laboratory during his postgraduate studies in Cambridge. (photograph courtesy of Jane Lamb.)




were no gene-specific probes available and it was necessary to use complex, time-consum-ing methods to separate newly formed proteins from those existing before the induction of the defence response. after Dixon moved to royal holloway College, University of London, in 1978, there were constant visits between egham and oxford, where Chris had become a Departmental Demonstrator in biochemistry (with his own laboratory, as a browne research Fellow at Queen’s College). the collaboration between Lamb and Dixon was both long-last-ing and productive; by the time of Chris’s death it had yielded 114 joint publications, the first being in FEBS Letters in 1978 (2) and the last in EMBO Journal in 2004 (39).

the step from studying wound-induced to pathogen-induced defence responses was a seminal one. throughout the rest of his life Chris focused on plant–pathogen interactions and defence signalling. although he had an enormous breadth of understanding and published on a very broad range of specific topics, opening up huge areas of research, everything he did was intimately related to his primary interest of plant disease signalling.

During this period, Chris and his student mike Lawton visited Klaus hahlbrock’s labora-tory at the University of Freiburg, to learn techniques for in vitro translation of rna (as a method for assessing whether new genes were being switched on) and applying it to the bean cell culture system. by an interesting turn of fate, hahlbrock was an adviser (non-resident Fellow) for the salk institute in La Jolla, California. this prestigious biomedical research insti-tute was then contemplating the establishment of a new plant biology programme; hahlbrock recognized Chris’s leadership potential and recommended him for the director’s position, an action that showed remarkable foresight because Chris had not yet held any permanent faculty position. Chris was appointed, moved in 1982, and rapidly built up one of world’s major plant biology programmes.

la Jolla

in early 1983 Chris moved his family to southern California and settled into a neighbourhood called University City. During the following decade and a half, Chris and Jane sent Catherine and William to Californian schools. William remains in California to this day. Despite family adjustments, this period was a remarkably productive one for Chris’s laboratory. During his 16 years at the salk institute, the lab underwent a transformation from a small laboratory with a few phD students to a large and very productive postdoc-driven lab of more than 20 members (figure 2). it quickly became one of the world’s leading laboratories, not only in plant pathol-ogy but also in several areas of modern plant biology, including cell cycle control (16, 24, 27), the control of secondary metabolism genes (see for example, (5–14, 17–19)), and signalling through protein kinases (21, 22).

this transformation was not always painless. as beat Keller, a postdoctoral worker from the mid 1980s, explains:

the research group of Chris Lamb was among the first to use molecular biological approaches to study plant-specific metabolism, in particular defense reactions induced by pathogen elicitors (see, for example, (3)). Consequently, the lab of Chris Lamb became a hotspot … attracting a large number of postdocs from all over the world, but in particular from europe. most of them had their own fellowships, and the number of people in the lab grew rapidly. postdocs profited from Chris Lamb’s mentoring, but also from the freedom he gave the people in his lab to develop their own research directions. this influx of people both from classical phytopathology and plant physiology,




as well as from molecular biology of microbial or animal systems, created an extremely creative and stimulating research environment. the integration of new tools such as plant transformation (in those days done with tobacco) and pCr [polymerase chain reaction] revolutionized the way biological problems could be approached in plants, and the study of plant gene expression became a hot topic. the new developments were sometimes faster than the adaptation of infrastructure to the new needs: when the first tobacco plants came out of tissue culture, there were no growth room facilities or greenhouses available to pot and grow them. this challenge was solved by a makeshift greenhouse frantically built by the facilities department of the salk institute.

one of us (J.C.) remembers at least one windy day when the ‘greenhouse’ was uplifted from its stakes and began to blow towards the ocean. Luckily, Yonaton elkind was able to snatch up the plastic, and, although the plants blew over, they survived. the situation was rectified slowly over the next decade with the hiring of new faculty.

the plant growth facilities were certainly primitive, yet the pace of science in Chris’s lab was nothing but remarkable. roger pennell remembers:

soon after moving to salk, i noticed Chris complaining that his publications were appearing in the journals at a rate of fewer than one a month. Chris was incredulous. his lab was large and produc-tive, at the center of the mechanism of plant disease resistance, and, well, one paper a month just wasn’t going to cut it. investigation revealed that the rate-limiting step was the process by which data were committed to text. a day or two later, Chris was talking into a tape recorder. slumped in his chair in his office, looking out of a window, he was muttering with typical intonation and at typical speed into a microphone. this went on all day, and then for several days. When he was finished, and under pressure to explain, he said he had been writing papers! to get his publications back up to speed—to save on time—Chris had taken to using dictation! Completed tapes were levered out, transcribed by his long-term administrative assistant, Cindy Doane, and returned for

Figure 2. Chris (second from the left) at the salk institute with (from the left) peter Doerner, Detlef Weigel Formemrs and Joanne Chory.




proofing. this took the form not of massive re-structuring, deleting, inserting, re-wording, re-emphasizing, de-emphasizing, re-planning and crumpling up and hurling towards the trash bucket, but of the one-off addition of occasional proofreading marks—a ‘delete and close the gap’ here, a ‘no paragraph: keep sentences together’ there—in the margins of a handful of the pages. one-paper-a-month turned into one-a-week. order was restored. this anecdote speaks to something that was important about him—his very special way of seeing, distilling, and packaging a piece of science, especially a piece of science that goes on to get cited over and over again.

after demonstrating how regulatory regions in several plant defence genes were responsi-ble for coordinated gene activation during induced chemical defences against pathogen attack, Des bradley and others went on to show that an ultra-fast hardening of plant cell walls, via the cross-linking of cell wall proteins (15), created a barrier that is the first line of defence against a fungal pathogen. seeking a trigger for this cross-linking, Chris’s team found that hydrogen peroxide was involved (26). they showed that hydrogen peroxide was not simply a cross-linking or potentially antimicrobial agent; rather, it also acted as a signal, not only for local disease resistance in the infected area but also at a distance from the infection site in the uninfected leaves of the plant (28–31). so-called ‘micro-oxidative bursts’ of hydrogen peroxide production were observed near the veins in uninfected leaves if a lower leaf of the plant had first been infected with a pathogen, and these were necessary for establishing immu-nity, or systemic acquired resistance (sar), to disease throughout the plant. Commonalities between induced resistance in plants and animals were illuminated by subsequent seminal discoveries of the involvement of nitric oxide (32, 36) and long-distance lipid signals (37) in plant defence. taken as a whole, this work provides the basis for a very simple and provoca-tive model for plant disease resistance that integrates an oxidative burst at the cell surface with transcription-dependent defence-gene activation and host cell death (34, 35, 38). this elegant series of studies well illustrates Chris’s approach as a scientist, and in particular his keen sense for the importance of the unexpected, initially puzzling result, his skill in designing experiments to illuminate their significance and mode of action, and his talent for (and delight in) incorporating new data into a larger, coherent intellectual framework. his discoveries have had a major impact on our thinking of how plants respond to microbial attack, and resulted in Chris’s recognition by the institute for scientific information as one of the most highly cited researchers in the plant and animal sciences.

one of the great pleasures of working alongside Chris was that he really enjoyed work-ing out ideas and discussing hypotheses. as recounted in several stories of mike Lawton and rick Dixon, Chris created an immensely enjoyable yet scientifically rigorous environment in which students and co-workers could sharpen their own critical thinking skills and develop as independent scientists. these discussions were not only in the lab or his office. in California, the oxford pubs were replaced by Friday evening ‘lab meetings’ at a mexican bar/restaurant. Chris had also discovered a hole-in-the-wall indian restaurant, which served one of the hottest lamb vindaloos on the planet, and visits there were both memorable and painful.

Chris was a tireless traveller during his tenure at the salk institute, and never seemed to suffer from jet lag. he must have been one of United airlines’ most frequent flyers, with a dizzying schedule of trips to asia, europe and south america. he loved recalling his adven-tures on these trips, from being stranded for days in far-off places to the time, on his first trip to Venezuela, when he thought he had been kidnapped by armed bandits (the security guards for the wealthy person he was visiting on behalf of the salk’s fundraising efforts). even in his absence, the lab continued to run productively. this was partly due to the ease of communi-




cating with the lab via the internet, and also to senior scientists in the lab, including michael Lawton (who returned for a second postdoctoral period) and peter Doerner. susie hedrick, Chris’s long-time lab manager, kept the lab running efficiently, while also performing experi-ments.

throughout these years, Chris was an actively engaged member of the salk community, serving on multiple committees and gracing them with his uncanny ability to synthesize the musings of those around him and turn them into concrete action plans. the salk’s commit-ment to excellence had a huge influence on Chris’s thinking when he later found himself in a position of leadership at the John innes Centre (JiC) (see below).

Chris’s legacy in La Jolla is not just his own science but also the scores of postdoctoral workers he mentored who now run their own laboratories and continue his legacy—these include Carole Cramer, steve Clouse, mike Lawton, mona mehdy, roger pennell, beat Keller, Jan Kooter, massimo Delledone, peter Doerner, michel Dron, Ken shirazu, alex Levine, Didier reinhardt, norbert sauer and Gary Loake. massimo Delledonne remembers Chris with particular affection because not only was he his mentor, he also found funding for him on three occasions when he was unexpectedly left without fellowship support, after hav-ing travelled to the Usa or edinburgh from his home in italy.

During this period, Chris also identified and mentored young faculty who have made La Jolla one of the top places for plant science in the Usa. Chris realized early on the power of Arabidopsis, recruiting J.C. in 1988, before she had published a single paper on Arabidopsis. Detlef Weigel was recruited in 1993 by Chris and J.C., to round out the original plant biology crew. Chris also had a role in recruiting plant faculty to the University of California at san Diego (UCsD) and the scripps research institute. as steve Kay, the current Dean of biology at UCsD, put it:

in addition to his science acumen, Chris’s generosity is what i will remember most. i owe a great deal to his counsel, especially as i pondered taking the dive from the east Coast to La Jolla. i recall that Chris encouraged me not to think of san Diego, as he put it, as ‘the dumb blonde of american cities’, but to consider how wonderful the mesa was as a place to grow professionally. i was concerned about collegiality, and Chris assured me that i would end up with more colleagues and friends ‘than a sheep has ticks’. he became a very good colleague and friend, and i owe much to his support and guidance, especially when there were difficult decisions to be made. i will miss him terribly, but there is much of Chris’s influence woven into the fiber of our plant biology com-munity here—a community he began and never stopped fostering.

today, La Jolla boasts 16 principal investigators at the salk and UCsD, seven of whom have been elected by their peers to the Us national academy of sciences (steve briggs, J.C., maarten Chrispeels, Joe ecker, mark estelle, steve Kay and marty Yanofsky (and Detlef Weigel, who has since moved to Germany)). La Jolla plant scientists have a deep footprint in the scientific literature, having published more than 1500 primary publications, which have been cited in the literature close to 100 000 times.

in addition to founding and nurturing the plant biology Laboratory at the salk, Chris was also instrumental in facilitating the early development of another programme that was soon to join salk among the premier places for plant science research in the Usa. the president of the salk institute was a friend of a member of the board of the ardmore, oklahoma-based samuel roberts noble Foundation, and the Foundation had provided a significant five-year start-up grant for the new plant biology programme at the salk. Chris saw the potential for also building a world-class programme at noble, fed by significant interactions between the




two organizations, and was instrumental in making that happen. after the establishment of noble’s plant biology Division in 1988, Chris was a constant visitor to ardmore. the noble Foundation renewed its five-year support for Chris’s programme, which now featured a joint postdoctoral fellowship programme, whereby selected fellows would spend 18 months in ardmore and 18 months in san Diego. this was critical for nurturing the noble plant biol-ogy programme through its formative years. between 1989 and 1999, 15 postdoctoral fellows worked on this programme, during a period that saw more than 80 joint publications from the two institutes. the interactions were facilitated by yearly retreats, alternately in California and oklahoma/texas, featuring salk and noble faculty, postdoctoral workers, and students, and an array of high-profile guest speakers. many of these noble/salk postdoctoral fellows now hold senior positions at prestigious universities and research institutes throughout the world.

Chris loved the american lifestyle. he often spoke of its endless optimism, which was very much a part of his character. however, he remained quintessentially british at heart, and finally made the move back across the atlantic in late 1998, briefly holding the regius Chair in plant science at the University of edinburgh. a greater challenge soon became available, and Chris moved to the JiC as its director later that year. he was determined to make a mark on UK plant science, and slowly but surely reorganized the science at the JiC, focusing on ‘excellence and relevance’, as well as ‘transparency’, implementing the high standards he had learned at the salk.

edinBurgh

For several years before he moved to edinburgh in late 1998, Chris had been following adver-tisements for vacant chairs in the UK, and shortly before his mother passed away in the sum-mer of 1998 he felt a strong responsibility for the wellbeing of his parents and wanted to be close. at about this time, the regius Chair in botany at edinburgh University was advertised. Chris was interested in this position because he perceived plant sciences at edinburgh to have exciting potential as a blend between recently recruited younger lecturers and well-established renowned professors, embedded within the famous institute for molecular biology and with links to a world-class botanical garden. he also expressed a desire to make a contribution to science in britain.

although Chris stayed in edinburgh for only nine months, his contributions and leader-ship were impressive. plant sciences was housed in the Daniel rutherford building, which had been inaugurated in 1965. peter Doerner recalls that Chris recognized immediately that a concerted effort to improve the facilities was also a great way to stimulate a discussion among the academic staff about the coherence and future direction of the department. he delegated the writing of sections of a proposal for Joint infrastructure Funding to many of the staff, but he also laid out a coherent vision, focused on signal transduction and plant development, for the bid. Ultimately, this led to a successful refurbishment of the Daniel rutherford building, completed in 2003. Chris was also instrumental in attracting funds (from the Leverhulme trust and akkadix, a Californian plant biotech company) to set up growth-room facilities and to conduct a large-scale plant genomics project, in which most of the junior faculty were involved, thereby enhancing both communication and facilities within the department.




norwiCh

shortly after Chris’s arrival in edinburgh, the post of director of the JiC, in norwich, was advertised. Chris was appointed to the position in 1999. by then, he had published 180 papers and had established an enviable citation record—he had an h-index (a measure of scientific productivity and impact) of 68 and had published five papers in Cell (4, 15, 20, 25, 31) with a citation average of about 650, one of which (25) had a citation index of 1380. his review ‘the oxidative burst in plant disease resistance’ (29) remains the second most highly cited paper ever published in Annual Review of Plant Physiology and Plant Molecular Biology. the demands of leading and managing a research centre of some 850 people, however, inevitably led to a marked decrease in his front-line research effort. the size of his research group, and the commitment of his time to research, diminished to such an extent that he managed to devote only about 10% of his time to this activity. he was necessarily reliant on the help of others for the day-to-day running of his group but he remained intellectually engaged in the programmes and in the career development opportunities for his team. over the period 1999–2009 the team included natalie buhot, matthieu Chabannes, Gary Creissen, Casey Crooks, Jun Fan, anita maldonado, Kanu patel, helen reynolds, maria romero-puertas, Christine rusterucci and Jurgen Zeier.

although he was not as productive in terms of science output as in the late 1980s and 1990s, he sustained his interest in fundamental areas of plant defence and continued to publish in top journals (including Nature, Science, EMBO Journal and Proceedings of the National Academy of Sciences of the USA), so retaining his remarkable ability to present his work in a way that ensured maximum impact. taking advantage of the JiC environment and its wide range of knowledge and expertise, he worked to integrate studies of the mechanisms of signal transduction with areas of more general relevance, for example non-host and basal resistance, natural genetic variation for resistance, biotic/abiotic signal cross-talk and the costs of resist-ance in crops.

Science researchthe Lamb group’s research focus at the JiC was based on two technical approaches (gain-of-function mutation and loss-of-function mutation), combined with the ectopic expression of specific genes, to analyse signalling in plant–microbe interactions. activation tagging with t-Dna that harboured cauliflower mosaic virus 35s promoter enhancer elements was used in the gain-of-function approach to identify two important Arabidopsis thaliana genes CDR1 and CDS2.

Aspartic protease and peptide elicitorsan activation-tagged Arabidopsis line (constitutive disease resistance 1-Dominant, CDR1-D) that overexpressed an aspartic protease was shown to be resistant to virulent strains of Pseudomonas syringae, an important plant pathogen that causes disease in a wide range of plants, including wheat, barley, legumes, brassicas and trees (40). Uninfected CDR1-D plants developed salicylic acid-dependent micro-lesions, reminiscent of the hypersensitive response induced by systemic micro-oxidative bursts in the establishment of sar, and exhibited elevated expression of defence genes such as those encoding pathogenesis-related proteins pr1 and pr2 and rbohF, the subunit of the naDph oxidase that contributes to the ‘oxidative burst’ (see above).




the CDR1 gene and complementary Dna were cloned and used for overexpression and antisense downregulation of CDR1 in Arabidopsis. these experiments confirmed that CDR1 functions in disease resistance. in addition, micro-grafting experiments showed that CDR1 activation leads to the systemic induction of defence responses, and site-directed mutagenesis confirmed that protease activity was required for this function (39). aspartic proteases have been implicated in the regulation of a variety of biological processes, and the CDr1 protein could function in response to the presence of a virulent pathogen in several ways. one attrac-tive possibility is that CDR1 activation leads to the generation of an extracellular peptide elici-tor that could function as a mobile signal, in a similar fashion to the wound response mediator systemin (pearce et al. 1991).

Abscisic acid and the interrelation between abiotic and biotic stress responsesin a search for genetic components in salicylic acid-mediated signalling mechanisms, salicylic acid-deficient (NahG) Arabidopsis plants were screened for complementing functions by acti-vation tagging (44). the screen involved inoculation with virulent P. syringae pv. maculicola (Psm) chromosomally tagged with a luciferase operon to allow rapid quantification of bac-terial growth in planta (42). no mutants with enhanced resistance to virulent bacteria were recovered, but one line (constitutive disease susceptibility2-1D (cds2-1D)) was identified that supported increased bacterial growth and was compromised both in basal resistance triggered by microbe-associated molecular patterns and in sar. molecular characterization of the mutant showed activation of 9-cis-epoxycarotenoid dioxygenase5 (NCED5), a member of the NCED gene family, which catalyses a key step in the biosynthesis of the plant growth and abiotic stress response regulator abscisic acid (aba).

the cds2-1D plant has elevated levels of endogenous aba. by crossing cds2-1D with mutants that carry genetic defects in the aba biosynthetic pathway, it was possible to show (44) that aba suppresses resistance to the hemibiotrophic bacterial pathogen P. syringae. extension of the study to other pathogens, including the biotrophic oomycete Hyaloperonospora arabidopsis and the necrotrophic fungus Alternaria brassicicola, sug-gested that aba accumulation compromised resistance to the former but promoted resistance to the latter. analysis of salicylic acid, jasmonic acid and aba in wild-type, cds2-1D and aba-deficient plants challenged with P. syringae showed that aba promotes jasmonate accumulation and has a complex, antagonistic relationship with salicylic acid. this was an interesting extension of seminal work done by Chris and others many years previously (33) that showed an inverse relationship between systemic resistance of plants to microorganisms and to insect herbivory, correlated with an inverse relationship between the levels of jasmonate and salicylic acid in challenged plants. the analysis of cds2-1D provided genetic evidence for cross-talk between pathogen infection and abiotic stress and suggested that aba modulates diverse plant–pathogen interactions partly by balancing jasmonate-based and salicylate-based biotic stress signal pathways.

Lipid transfer protein and mobile signalsscreening for mutants in sar has always been notoriously difficult, partly because sar is strongly affected by developmental stage and environmental conditions (Kus et al. 2002). Despite this, maldonado et al. (37) screened 11 000 t-Dna-tagged (loss-of-function) A. thaliana lines and found one, defective in induced resistance (dir1-1), that was com-promised in sar. although dir1-1 showed a wild-type local defence response, in terms




of oxidative burst, accumulation of salicylic acid and resistance to avirulent and virulent P. syringae, the expression of pathogenesis-related genes was abolished in non-inoculated distant leaves, and dir1-1 failed to develop sar to virulent P. syringae or to Peronospora parasitica. petiole exudate experiments suggested that dir1-1 is defective in the production or transmission of an essential mobile signal from the infected leaf; DIR1 was shown not to be required for salicylic acid accumulation (which is required for sar), but it remained possible that salicylic acid might contribute to the sar signalling pathway downstream of DIR1. DIR1 encodes a protein of 102 amino acid residues that has 53% sequence similarity to a Phaseolus vulgaris non-specific lipid transfer protein (Ltp); the presence of a hydro-phobic amino-terminal signal sequence suggested that Dir was apoplastic. the possibility that Dir1/Ltp itself might be a mobile signal was discounted because overexpression of DIR1 was not sufficient to induce sar, and the possibility remains that Dir1/Ltp could act as a translocator for the release of a mobile signal and/or could chaperone the signal through the plant. subsequent crystallization and structural analysis of the protein from A. thaliana (41, 43) showed that Dir1, which has some specific features that define it as a new type of plant Ltp, is able to bind, with high affinity, two monoacylated phospholipids side by side in an internal tunnel. a putative orthologue of At-Dir1 has been detected in the phloem sap of tomato plants (mitton et al. 2008), supporting the notion of common sar signalling molecules in different species.

Science managementRestructuringWithin a year of his arrival, partly as a consequence of the financial situation and partly through a perceived need to focus on and create headroom for new areas of science, Chris took the JiC through a process of science restructuring and realignment. starting from the original ten departments he created six new ones that better reflected the biotechnology and biological sciences research Council (bbsrC) grants panels and simplified reporting to the institutes’ assessment exercise (the institutes’ equivalent of the universities’ research assessment exercise). the departmental restructuring was not to everybody’s liking—in particular, many of those who had been part of the Department of Genetics were understandably dismayed to see the department disappear from the JiC, whose first director (William bateson Frs) had coined the term ‘genetics’. however, a beneficial consequence was that people suddenly found themselves among new departmental colleagues, with the result that new scientific alliances were forged without losing the previous associations, resulting in greater communication and interaction across the site. the restructuring also provided a means of integrating the members of the former nitrogen Fixation Laboratory, which had moved to norwich from sussex a few years earlier, with the formation of the departments of biological Chemistry and molecular microbiology. Chris recognized the strengths in these departments and for some years his laboratory was located in the same building, providing a point of connection between the JiC’s microbiological and plant research communities.

Grant proposalsin 1999 the JiC was not eligible to apply for so-called ‘responsive mode’ grants from its core sponsor, the bbsrC. one of Chris Lamb’s early achievements at the JiC, together with other senior JiC staff, and in particular rico Coen Frs, was to bring this to the attention of the bbsrC, as a consequence of which bbsrC granting policy was changed and institutes




became eligible to apply for responsive mode grants, with one proviso: there would be a cap on the amount of funding for which a given institute could apply within a given year. to ensure that every application stood the best possible chance of success, Chris set up internal panels to assess and improve proposals before submission. this represented a transformation in behaviour and had considerable potential for friction, but because it resulted in a dispro-portionately high success rate for the JiC, project leaders were quick to accept that the system improved their proposals, and those on the internal panels gained a much better knowledge of the activities and aspirations of the JiC in general. Chris had put in place a mechanism for achieving excellence but had also achieved a significant degree of improvement in scientific communication across the site. the policy led the way beyond the JiC: other institutes and universities have since followed the same path.

Appointments and tenure track careersChris’s mantra was ‘excellence and relevance’. his definition of relevance, in the context of the JiC, encompassed food security, developing bio-energy, delivering sustainable agricul-ture, improving healthy ageing and discovering new therapeutic agents and targets. to attain relevance he established what he called ‘clusters’ of a range of talents, to translate fundamen-tal discoveries into practical outputs. he ensured excellence by appointing the best people to project leader positions and one of the main changes that he brought about was to have non-departmental quality control before appointments were made. the whole of the JiC was involved in appointments, as candidates were seen by representatives from each department, interviewed by a panel drawn from across the JiC (and including an external assessor) and assessed by an interdepartmental appointments committee. Chris also took the point of view that, having appointed the best, they should be given a level of support that would enable them to excel—this in turn implied ring-fencing of a significant level of recurrent resources and demanded that the appointments and subsequent mentoring processes be robust. he therefore put in place a tenure track system whereby project leader appointments were for an initial period of five years, with the appointee being carefully mentored, and reviewed around the three-year or four-year mark. this gave the appointee sufficient time to establish a grants portfolio, a moderate size of research group and a good publication record, but also provided at least a year to seek another position should they not gain tenure. it also ensured that new faculty were given proper support and mentoring, and that excellent scientists were appointed in the long term. such a system represented a massive change in the JiC appointments pro-cedure but simply reflected Chris’s desire to promote excellence.

StudentshipsChris recognized the important role of the training of young scientists in the remit of the JiC, and he launched several initiatives designed to enhance studentship programmes and to empower the student community. he provided students with their own budget that ena-bled them to organize social events and a well-endowed seminar series entitled ‘science in society’. in addition, a more student-centred approach began to develop within the work-ings of the Graduate studies Committee, which helped to raise the standards of training and supervision by sharing good practice across the site. With the support of the John innes Foundation and the associated sainsbury Laboratory, Chris instituted a flagship four-year rotation phD programme, designed to attract the very best students from the UK and around the world. although the Wellcome trust had pioneered this form of phD studentship in the




UK, the JiC programme was the first to be established in the field of plant and microbial sciences. a very high academic standard was set for recruitment, and successful candidates received a significantly enhanced stipend to reflect the prestige attached to the programme. During their first year, each rotation student gained experience of research in three different laboratories in different departments before selecting a research topic and a supervisor for their thesis work. the rotation phD programme had a profound effect on both the qual-ity of student training at the JiC and the quality of research. it also helped to develop new interdisciplinary research collaborations between project leaders from both the JiC and the sainsbury Laboratory. this networking feature was clearly beneficial for both the students and the host institute and moreover proved highly successful in attracting external funding for the sponsorship of phD programmes at the JiC, most notably from the european Union’s marie Curie early stage training and the bbsrC. there can be little doubt that one of Chris’s many legacies has been to leave a much more vibrant and robust phD programme at the JiC.

Computational biology and bioinformaticsWhen Chris joined the JiC there was relatively little in the way of computational biology in its broad, interdisciplinary sense. Chris recognized that the new ‘omics’ technologies flourishing at the JiC in the early 2000s would require extensive bioinformatics support. he organized a one-day discussion meeting, bringing leading bioinformaticists to the JiC, including Lincoln stein (Cold spring harbor Laboratory), ewan birney (sanger), simon Lovell (Cambridge), alan robinson (european bioinformatics institute) and simon hubbard (University of manchester institute of science and technology). this meeting strongly supported the creation of a Computational biology Unit at the JiC. a panel of external experts, chaired by professor Janet thornton Frs, was subsequently invited to provide advice on how computa-tional biology/bioinformatics could best be integrated into JiC science. the panel found that there was an immediate need to increase critical mass in bioinformatics to increase support for databases, software and ‘omics’ technologies. their specific recommendations were, however, problematic in that they suggested the appointment of about 20 new staff. Chris was strongly in favour of expanding bioinformatics and embedding it within departments, but with limited resources available not all the recommendations of the panel could be realized. he therefore established a procedure whereby those groups who required computational biol-ogy in their research included a request for it in grant applications, and successes in grants were augmented from the JiC’s central resources, leading to the creation of three new posts in bioinformatics. the next challenge was the emergence of systems biology as a major new impetus in the biosciences. the JiC had the necessary infrastructure and scientific breadth to make a strong contribution to systems biology, and many interdisciplinary collaborations had been established. in-house expertise in physics and mathematics was, however, lacking at the JiC, so there were limited opportunities for accurate modelling and prediction of the diverse plant and microbial systems studied there. Chris realized that new project leader appointments in computational biology or mathematics would be necessary to fill this gap. over a period of five years he appointed four project leaders with expertise in mathematical modelling, to complement the Centre’s existing strengths in statistical modelling, comparative genomics and macromolecular modelling. to support the new cadre of modellers, Chris created a new Department of Computational and systems biology that integrated this biomathematical and biophysical research with the bioinformatics group he had built up previously. this nucleus




of modellers at the JiC has driven many exciting collaborations that integrate theoretical and experimental approaches, establishing the JiC as an international centre of excellence in mechanistic modelling of plants and microbes.

CommunicationChris was a paradox when it came to communication. on the one hand he was a great friend to, and an effective communicator with, those whom he liked, respected and trusted, but on the other he could be terse when confronted with people who failed to communicate their positions in an intellectually robust manner. there were those, especially those who were scientifically close to him, who thought that he was a paradigm for scientific communication—his lectures have been described by mike Lawton as ‘watching a fine tapestry being woven in front of your eyes’. at a personal level, when at ease with his peers in his field of study, he was witty and charming company; but he was also inherently shy and this could easily be interpreted as arrogance—he has been described as ‘somewhat inscrutable’ by those who didn’t know him well. this shyness could lead people to assume that he was overly serious, but in fact he had a well-developed sense of humour that can be best appreciated from an e-mail that he sent from a bbsrC management Development programme at roffey park (figure 3) in which he pretended to be imprisoned against his will and described his fellow ‘inmates’ and his attempts to escape. the style illustrates a witty side of Chris’s character that was not always generally appreciated on a day-to-day basis.

in some ways, Chris was not at ease in public speaking roles, but he undertook them because he believed in, and was a passionate advocate for, what he saw as a remarkable area of science. When talking about plant science, Chris exuded enthusiasm. he talked as if all of his audience shared his enthusiasm and understanding—those who were non-scientists found the first to be infectious and the second flattering. stuart holmes, a non-scientific member of the JiC’s governing council and the chair of council for the University of east anglia (Uea), recalls that Chris never tried to blind him with science but treated his contribution to the JiC on level terms, which led to a deep and abiding mutual trust.

Chris was a very strong supporter of the JiC Friday seminar programme—despite the fact that he could never manage to arrive before they started—and he not only empowered the organizing committee to ensure that JiC staff were exposed to the very best in world science by attracting world-class speakers but he also encouraged younger members of staff to intro-duce the seminars and meet the speakers.

Chris recognized the vital importance of explaining research to the wider world and ensur-ing that discussion about its scientific and moral dimensions is well founded. he was well aware of the politics of science, and in particular the issues surrounding food security and genetically modified plants. Faced with the realities of ill-informed and prejudiced campaign-ing, he used the influence of his office as JiC director to try and ensure that public debate and consideration of the issues was informed, balanced and proportionate. one means he chose for doing this was a series of dinners in the house of Commons, formally hosted by Charles Clarke mp, in whose constituency the JiC was located. each dinner was given a different theme, for example food security, sustainable agriculture, developing bio-energy, or healthy ageing. Chris usually arrived carrying his folding bike, on which he had cycled from Liverpool street station, prompting chaos as the house of Commons security staff insisted that it be put through the X-ray machine. the dinners brought together about 15 people, some of whom were John innes scientists but most of whom were leading scientists from elsewhere, leading




politicians (of all parties), civil servants, journalists, and representatives of non-governmental and campaigning organizations. Chris very deliberately included some of the best-known peo-ple in the country on all sides of the debate. Contested argument was at the core, and so the discussions were vigorous but courteous. Chris provoked, using his own brand of charm and humour, and everyone always left in a good mood, having enjoyed stimulating exchanges. the

Figure 3. email from Chris Lamb to tony maxwell and anne osbourn. in 2007 the bbsrC encouraged its staff to attend a management Development programme held at roffey park near Gatwick. maxwell and osbourn were the first attendees; Chris and other senior staff attended a later course. Chris wrote this email to tony and anne in the style of the situation comedies Dad’s Army and It Ain’t Half Hot Mum; it contains many cryptic refer-ences to bbsrC personnel and institutes. the annotations were made by tony maxwell.




dinners did not seek collective conclusions, but they did try to ensure that everyone understood the others’ positions and reduced consideration of the issues to their scientific basis. the proc-ess was a testament to Chris’s faith in the power of argument and the scientific method, and it had a very real impact on the tone and nature of this important public debate.

Chris’s love of middlesbrough Football Club took him to most of the english premiership stadia, where he thoroughly enjoyed discussing matches and players and brought his scien-tist’s logic to bear on a sharp analysis of tactics. he was, as ian Gibson mp described him, a man for all seasons—able to communicate as easily with government ministers and members of parliament as he was with the football fan next to him in the pub, or with a student at the beginning of their career. this ability to communicate and interact at all levels was exem-plified by an incident that took place in edinburgh, just after Chris had been announced as the new JiC director but before he was in post. the event was a public outreach play at the edinburgh Festival, exploring the issues surrounding genetically modified food. one of the on-stage ‘experts’ (the inverted commas are hers), there to answer questions from the audience about crop biotechnology at the end of the play, was belinda Clarke, a young JiC phD student, who was shocked to spot her new director in the audience. however, she need not have wor-ried because she found that he was supportive of her to the point at which they became an impromptu double act in the face of some robust views against genetic modification, despite never having met before. Chris took a clear and intellectually rigorous stand on genetic modi-fication, championing not simply the science but also the morality of a scientifically based approach that would help to meet the global food challenge and other health issues. Chris commented afterwards that they were ‘digging for potatoes’, without clarifying whether this was a criticism or an endorsement—simultaneously communicative and inscrutable.

this unusual use of phrases and idioms typified Chris; he enjoyed language and there were several occasions when people would leave a meeting with him and head straight for the dictionary. after a while at the JiC, his love of obscure management-speak and metaphor spawned ‘Lambword bingo’, a variant of buzzword bingo. alison smith, head of metabolic biology at the JiC, believes that he almost certainly was well aware of this, and probably played to it at times. ‘Digging where the potatoes are’, ‘getting the ducks in a row’, ‘less is more’ and ‘do good things and good things will follow’ were favourite but rather obvious phrases of his; more obscure ones included ‘staring down the well’, ‘coming in on the mez-zanine level’ and references to the JiC’s role in helping people ‘not to make better widgets, but to better make better widgets’.

Distractionsas with any new director, Chris inherited several issues that were not of his making, one of which—the syngenta research agreement—proved to be an unwelcome distraction. the JiC entered into an alliance with Zeneca plc in 1998, with the purpose of ensuring that discoveries from the fundamental research on wheat at the JiC were used to develop new wheat varieties (figure 4). on 16 July 2001 the foundation stone was laid of a £13 million state-of-the-art Genome Centre that was to house separate JiC and syngenta (as Zeneca had become) labo-ratories on the ground floor and a new venture, the norwich bioincubator, partly funded by the east of england Development agency, on the top floor. Within little more than a year, however, syngenta had signalled its intention to withdraw from the alliance, although the bioincubator flourished. although the decision had a minimal, and short-term, impact on JiC research, the withdrawal had a significant effect on JiC finances; the details of the termination




and withdrawal were complex and the consequent discussions and negotiations were delicate, extensive and a distraction that Chris would not have contemplated when he accepted the post of director.

a second major distraction, which arose during Chris’s directorship, was that of govern-ance. the issue was prompted by disquiet, elsewhere, about the nature and appropriateness of the governance of research Council institutes, including the JiC. this was particularly irksome as far as Chris was concerned because he was very happy with the composition and behaviour of his governing council (chaired by sir John beringer for much of Chris’s tenure as director), greatly valued the support it gave him and saw no reason for appreciable change. What began as a light-touch review of the governance of research Council institutes, performed over the period July 2005 to January 2006, evolved into a much larger issue and became a major, time-consuming distraction for Chris that was still unresolved at the time of his death.

Further significant claims on Chris’s time came in the shape of perturbations to the JiC’s financial health, changes in employment law (and in particular the european Union fixed-term work Directive), and the merger of JiC administration and support services with those of the neighbouring institute of Food research (iFr). the last task was delegated largely to a small group, but Chris was a prime mover in initiating the process; the success of the exercise, in terms of savings and efficiencies, are testimony to his determination and that of David White, the iFr director at the time. Chris firmly believed that excellent science requires equally excel-lent support and looked to the new, merged, support services to provide such excellence.

Given Chris’s immense scientific capabilities, there were many who felt that he should never have been burdened with such distractions, but it was in Chris’s nature to be involved and to ensure that any outcomes benefited science. he was unaccustomed to the level of administrative formality and behaviour that he encountered at the JiC, which was government-

Figure 4. Chris among the wheat in a JiC glasshouse. (photograph courtesy of andrew Davis, JiC.) (online version in colour.)




sponsored and had a comprehensive and detailed staff code that covered all areas of activity. he obviously understood that such a code was driven by employment and other legislation, but at times he was exasperated by what he saw as limitations on his plans for the JiC. Despite these potential limitations, in all his dealings over administrative or operational matters Chris never relented on the primacy that he accorded to obtaining benefit for science, which was always foremost in any of the arguments he deployed to either counter or initiate change. in the view of ian Crute, who was director of rothamsted research at the time, this approach was perhaps most evident in the way he pursued policies for recruitment and retention of scientific staff at the JiC. his insistence on seeking and supporting strong performers wherever they were to be found did not always sit comfortably with the more process-driven position adopted by those charged with ensuring compliance with codes of employment practice, but it undoubtedly contributed enormously to the success of the JiC.

The lamB legaCy

one of Chris’s main legacies was the way in which the norwich research park (nrp) devel-oped during his time at the JiC. Chris strongly supported the nrp, chaired the boards of direc-tors of the two nrp limited companies—nrp science (nrp-s) and nrp enterprises—and saw the research park grow substantially, to eventually comprise the JiC, the sainsbury Laboratory, the iFr, Uea, the norwich bioincubator, plant biosciences Ltd, the bbsrC Genome analysis Centre and the norfolk and norwich University hospital. When the last of these opened, the ‘field to fork’ metaphor became a commitment to interdisciplinary work-ing that linked plant and microbial sciences, food, health and sustainability at the nrp. not long before Chris’s death, the bbsrC showed its commitment to the nrp by establishing the norwich science and innovation partnership, which had a programme board, of which Chris was a member, and a science strategy board, which he chaired. Chris was instrumental in engaging not just the other directors of nrp-s but also the wider stakeholders, such as local authorities, in the nrp-s strategy. his vision for the nrp in the broader UK and global contexts catalysed numerous positive developments for the research park—David eastwood, who was Vice-Chancellor of Uea for much of Chris’s time at the JiC, recalls that Chris saw clearly, and worked assiduously towards ensuring, that the nrp was considerably more than the sum of its individual parts, integrating the science and technology to solve problems in biosciences and environmental science.

Chris was keenly aware of the political significance, as well as the social and economic benefits, of commercial exploitation and was strongly supportive of the general enhancement of the JiC’s commercial activities, including the machinery to ensure that both inventors and the JiC should receive proper rewards. he was instrumental in pushing forward a large number of elements of the JiC’s intellectual property and recognized that scientists who are willing to take risks to commercialize their science in the form of ‘spin-out’ companies not only brought valuable business experience to the JiC but also, importantly, needed career support to allow them to undertake such projects.

With his clarity of thought, incisiveness and vision, Chris played a key role in shaping the thinking of people he met, encouraging them to think in a wider context and providing a subtle safety net to inspire confidence in their own decision-making abilities. Critically, he believed deeply that plant and microbial science can change the world for the better and he




made others—no matter how junior—feel that they, too, could make an important contribu-tion to understanding. this ability, to give people the support and confidence to achieve more than they thought they could, served Chris well at all stages of his career and remains one of his legacies.

Chris had the ability, underpinned by sheer hard work and dedication, to make what he did look easy. he was unremittingly positive and had a great talent for seeing opportunities where other people might see only problems. When experiments seemed to fail, he could rapidly construct a model that made intellectual and scientific sense, fitted with the data from the ‘failed’ experiment and suggested new avenues and new experiments. above all, through his own sheer love of science and his keen interest in biology, he showed others that scientific research could be fun, exciting and useful, and taught them his very special way of seeing, distilling and packaging a piece of science in a way that achieved maximum effect.

this enthusiasm was coupled with the ability to consider and appreciate the constructive comments and thoughts of others, regardless of their source. he was particularly interested in learning the opinions and ideas of newly arrived career-track project leaders, to whom he would lend as much of an ear as he would to more established members of the faculty. similarly, he generously shared his experience on how to become a leader within one’s field—it was mentoring without lecturing, and it was greatly valued. as Lars ostergaard, a JiC project leader who often went with Chris to watch football, has put it, Chris expected hard work, dedication and the highest level of ambition, but in return he gave his postdoctoral workers and career-track project leaders the best possible conditions in which to thrive.

rick Dixon recalls that although Chris knew exactly what he wanted to do, and how to get there, he was anxious that his friends and colleagues should feel that they were part of the plan and he was considerate of their concerns. having listened to a valid criticism, he was immediately prepared to accede to and accommodate the other point of view; there was never a ‘stick-in-the-mud’ approach to any of his thinking. he was always keen to ensure that his approach, and by consequence the approach of the JiC, was as broad-minded as possible. the vision that he held for the JiC was always under review and swift to adapt to the ever-chang-ing needs of society and the various funding bodies that have consistently supported the work there. by revamping the groupings of the research to meet these needs, he ensured that the JiC continued to be relevant and an important part of global plant and microbial research.

sir ben Gill, a member of the JiC’s governing council and a former president of the national Farmers’ Union, describes Chris as the central rock for the JiC, about which every-thing else revolved. in his own unique style he gave a comfort to all who knew him that the Centre was in safe hands. but much more than that, by the sheer inspiration that he exuded he was always able to recruit the world-class scientists who are so crucial to maintaining the status of the JiC as a leading plant and microbial science research institute. moreover, he had a skill for talent spotting and for finding ways to continue to stretch and reward those around him. he was remarkably generous to those who worked with him as students, postdoctoral fel-lows or visiting scientists, invariably agreeing that they could take away their projects, which they had initiated under his supervision, to their new jobs. this, above all, is his major legacy: the careers he nurtured and the scientists he identified and mentored, some of whom remain at the salk and the JiC and some of whom have gone on to be professors and directors of major research institutions throughout the world. that legacy is very wide and its significance will grow as the world faces up to the important challenges—food security, renewable bio-energy, healthy ageing and sustainable agriculture—to which he devoted his time and energy.




perhaps the most telling testimony to Chris’s capability and leadership, however, came in the shape of data, published in Times Higher Education on 8 april 2009, from a worldwide survey of those institutions in plant and animal science that had amassed 25 000 or more cita-tions over the period 1999–2009, exactly coinciding with Chris’s directorship of the JiC. the top institution, on the basis of the impact of its publications, was the JiC. it is hard to imagine a better reflection of Chris’s dedication to quality and impact.

Chris was a devoted family man and was delighted by the arrival of sadie, his first grand-child, in July 2009. it was a tragedy that he should have been taken away so suddenly, from heart failure at home, only seven weeks after this happy event in his life.

honours and awards

1971–72 Fitzwilliam College, University of Cambridge: Wallersteiner exhibitioner1972–75 Fitzwilliam College, University of Cambridge: senior scholar1975–77 University of oxford: iCi research Fellow1977–82 the Queen’s College, oxford: browne research Fellow1979 University of Freiburg: british Council travelling Fellow1980 embo short-term Fellow1983 mcKnight Foundation research scholar1987 herman Frasch Foundation award of the american Chemical society Distinguished Lecturer in biology, boyce-thompson institute, Cornell University1990 Fellow of the american association for the advancement of science1992 Gottlieb memorial Lecturer, University of illinois2001 member of the european molecular biology organization Woolhouse Lecturer, society of experimental biology2008 Fellow of the royal society2009 Commander of the british empire

aCknowledgemenTs

We are especially grateful to Jane Lamb for her recollections of Chris as a young man, and for providing the photo-graph for figure 1. We are very grateful to the following friends and colleagues who made helpful comments on, and contributions to, this memoir: mary anderson, sir John beringer, mike bevan, mervyn bibb, nick brewin, Keith Chater Frs, belinda Clarke, Charles Clarke, rico Coen Frs, Gary Creissen, ian Crute, Caroline Dean Frs, massimo Delledonne, ray Dixon Frs, rick Dixon, peter Doerner, David eastwood, Jun Fan, Jeni Fox, ian Gibson, sir ben Gill, stuart holmes, steve Kay, beat Keller, mike Lawton, andy maule, tony maxwell, michael mcarthur, mike merrick, anne osbourn, Lars ostergaard, roger pennell, Keith roberts, alison smith, John snape, simon turner and Detlef Weigel Formemrs.

the frontispiece photograph was taken in 2008 by andrew Davis, JiC, and is reproduced with permission. (online version in colour.)




referenCes To oTher auThors

Kus, J., sarkar, r., Zaton, K. & Cameron, r. K. 2002 age-related resistance in Arabidopsis thaliana is a development-ally induced response to Pseudomonas syringae. Plant Cell 14, 479–490.

mitton, F. m., pinedo, m. L. & de la Canal, L. 2008 phloem sap of tomato plants contains a Dir1 putative ortholog. J. Plant Physiol. 166, 543–547.

pearce, G., strydom, D., Johnson, s. & ryan, C. a. 1991 a polypeptide from tomato leaves induces wound-inducible proteinase inhibitor proteins. Science 253, 895–898.

BiBliography

the following publications are those referred to directly in the text. a full bibliography is available as electronic supplementary material at http://dx.doi.org/10.1098/rsbm.2010.0014 or via http://rsbm.royalsocietypublishing.org.

(1) 1976 (With p. h. rubery) inhibition of co-operative enzymes by substrate analogues: possible implications for the physiological significance of negative co-operativity illustrated by phenylalanine metabolism in higher plants. J. Theor. Biol. 60, 441–447.

(2) 1978 (With r. a. Dixon) stimulation of de novo synthesis of L-phenylalanine ammonia-lyase in cell suspen-sion cultures of Phaseolus vulgaris. FEBS Lett. 94, 277–280.

(3) 1985 (With C. L. Cramer, t. b. ryder & J. n. bell) rapid switching of plant gene expression by fungal elicitor. Science 227, 1240–1243.

(4) 1989 (With m. a. Lawton, m. Dron & r. a. Dixon) signals and transduction mechanisms for activation of plant defenses against microbial attack. Cell 56, 215–224.

(5) 1991 (With m. a. Lawton, s. m. Jenkins, m. Dron, J. m. Kooter, K. m. Kragh, m. J. harrison, L. Yu, L. tanguay & r. a. Dixon) silencer region of a chalcone synthase promoter contains multiple binding sites for sbF-1, a factor closely related to Gt-1. Plant Mol. Biol. 16, 235–249.

(6) (With e. r. blyden, p. W. Doerner & r. a. Dixon) sequence analysis of a chalcone isomerase cDna of Phaseolus vulgaris. Plant Mol. Biol. 16, 167–169.

(7) (With m. J. harrison, m. a. Lawton & r. a. Dixon) purification of a nuclear protein which binds to three elements within the silencer region of a bean chalcone synthase promoter. Proc. Natl Acad. Sci. USA 88, 2515–2519.

(8) (With Q. Zhu) isolation and characterization of a rice gene encoding a basic chitinase. Mol. Gen. Genet. 226, 289–296.

(9) (With m. J. harrison, a. D. Choudhary, i. Dubery & r. a. Dixon) Cis-elements and trans-acting fac-tors for the quantitative expression of a bean chalcone synthase gene promoter in electroporated alfalfa protoplasts. Plant Mol. Biol. 16, 877–890.

(10) (With G. J. Loake, a. D. Choudhary, m. J. harrison, m. mavandad & r. a. Dixon) phenylpropanoid pathway intermediates regulate transient expression of a chalcone synthase gene promoter. Plant Cell 3, 829–840.

(11) (With r. a. Dixon, a. D. Choudhary, m. J. harrison, b. a. stermer, L. Yu, s. m. Jenkins & m. a. Lawton) transcription factors and defence gene activation. in Biochemistry and molecular biology of plant–pathogen interactions (ed. C. J. smith), pp. 271–284. oxford: oxford science publications.

(12) (With r. a. Dixon) properties of plant defense gene promoters. in Advances in molecular genetics of plant–microbe interactions (ed. h. hennecke & D. p. s. Verma), vol. 1, pp. 367–372. Dordrecht: Kluwer academic publishers.

(13) 1992 (With e. m. herman) arabinogalactan-rich glycoproteins are localized on the cell surface and in intra-vacuolar multivesicular bodies. Plant Physiol. 98, 264–272.

(14) (With a. m. Leyva, X. Liang, J. pintor-toro & r. a. Dixon) Cis-element combinations determine phenylalanine ammonia-lyase gene tissue-specific expression patterns. Plant Cell 4, 263–271.




(15) (With D. J. bradley & p. Kjellbom) elicitor- and wound-induced oxidative cross-linking of a proline-rich plant cell wall structural protein: a novel, rapid plant defense response. Cell 70, 21–30.

(16) (With m. C. martinez, J.-e. Jorgensen, m. a. Lawton & p. W. Doerner) spatial pattern of cdc2 tran-scripts in relation to meristem activity and cell proliferation during plant development. Proc. Natl Acad. Sci. USA 89, 7360–7364.

(17) (With G. J. Loake, o. Faktor & r. a. Dixon) Combination of h-box (CCtaCC(n)7Ct) and G-box (CaCGtG) cis-elements is necessary for feedforward stimulation of a chalcone synthase promoter by the phenylpropanoid intermediate 4-coumaric acid. Proc. Natl Acad. Sci. USA 89, 9230–9234.

(18) 1993 (With Q. Zhu & p. W. Doerner) stress induction and developmental regulation of a rice chitinase pro-moter in transgenic tobacco. Plant J. 3, 203–212.

(19) (With J. a. arias & r. a. Dixon) Dissection of the functional architecture of a plant defense gene promoter using a homologous in vitro transcription initiation system. Plant Cell 5, 485–496.

(20) 1994 plant disease resistance genes in signal perception and transduction. Cell 76, 419–422.(21) (With s.-h. Zhang, m. a. Lawton & t. hunter) Atpk1, a novel ribosomal protein kinase gene from

Arabidopsis. i. isolation, characterization and expression. J. Biol. Chem. 269, 17586–17592.(22) (With s.-h. Zhang, m. a. broome, m. a. Lawton & t. hunter) Atpk1, a novel ribosomal protein kinase

gene from Arabidopsis. ii. Functional and biochemical analysis of the encoded protein. J. Biol. Chem. 269, 17593–17599.

(23) (With n. J. bate, J. orr, W. ni, p. W. Doerner, r. a. Dixon & Y. elkind) Quantitative relation-ship between phenylalanine ammonia-lyase levels and phenylpropanoid accumulation in transgenic tobacco identifies a rate determining step in natural product synthesis. Proc. Natl Acad. Sci. USA 91, 7608–7612.

(24) (With p. Vera & p. W. Doerner) Cell-cycle regulation of expression of the hydroxyproline-rich glyco-protein HRGPnt3 gene in lateral root initiation. Plant J. 6, 717–727.

(25) (With a. Levine, r. tenhaken & r. a. Dixon) h2o2 from the oxidative burst orchestrates the plant hypersensitive disease resistance response. Cell 79, 583–593.

(26) 1995 (With r. tenhaken, a. Levine, L. F. brisson & r. a. Dixon) Function of the oxidative burst in hyper-sensitive disease resistance. Proc. Natl Acad. Sci. USA 92, 4158–4163.

(27) (With p. W. Doerner, J.-e. Jorgensen, r. You & J. steppuhn) Control of root growth and development by cyclin expression. Nature 380, 520–523.

(28) (With m. e. alvarez) oxidative-burst mediated defense responses in plant disease resistance. in Oxidative stress and the molecular biology of antioxidant defenses (ed. J. scandalios), pp. 815–839. Cold spring harbor, nY: Cold spring harbor Laboratory press.

(29) 1997 (With r. a. Dixon) the oxidative burst in plant disease resistance. Annu. Rev. Plant Physiol. Plant Mol. Biol. 48, 251–275.

(30) (With Z.-J. Guo & r. a. Dixon) release and biological activity of diffusible signal compounds from elicited plant cells. J. Plant Physiol. 151, 699–710.

(31) 1998 (With m. e. alvarez, r. i. pennell, p.-J. meijer, a. ishikawa & r. a. Dixon) reactive oxygen interme-diates mediate a systemic signal network in the establishment of plant immunity. Cell 92, 773–784.

(32) (With m. Delledonne, Y. Xia & r. a. Dixon) nitric oxide signal functions in plant disease resistance. Nature 394, 585–588.

(33) 1999 (With G. W. Felton, J. L. bi, K. L. Korth, s. V. Wesley, m. C. mathews, J. b. murphy & r. a. Dixon) inverse relationship between systemic resistance of plants to microorganisms and to insect herbivory. Curr. Biol. 9, 317–320.

(34) (With V. K. rajasekhar & r. a. Dixon) early events in the signal pathway for the oxidative burst in soybean cells exposed to avirulent Pseudomonas syringae pv. glycinea. Plant Physiol. 120, 1137–1146.

(35) (With r. K. Cameron, n. L. paiva & r. a. Dixon) accumulation of salicylic acid and pr-1 gene transcripts in relation to the systemic acquired resistance (sar) response induced by Pseudomonas syringae pv. tomato in Arabidopsis. Physiol. Mol. Plant Pathol. 55, 121–130.

(36) 2001 (With m. Delledonne, a. marocco & J. Zeier) signal interactions between nitric oxide and reactive oxygen intermediates in the plant hypersensitive disease resistance response. Proc. Natl Acad. Sci. USA 98, 13454–13459.




(37) 2002 (With a. m. maldonado, p. Doerner, r. a. Dixon & r. C. Cameron) a putative lipid transfer protein involved in systemic resistance signalling in Arabidopsis. Nature 419, 399–403.

(38) 2003 (With a. Kachroo, Z. he, r. patkar, Q. Zhu, J. Zhong, D. Li, p. ronald & b. b. Chattoo) induction of h2o2 in transgenic rice leads to cell death and enhanced resistance to both bacterial and fungal pathogens. Transgenic Res. 12, 577–586.

(39) 2004 (With Y. Xia, h. suzuki, J. borevitz, J. blount, Z. Guo, K. patel & r. a. Dixon) an extracellular aspartic protease functions in Arabidopsis disease resistance signaling. EMBO J. 23, 980–988.

(40) (With h. suzuki, Y. Xia, r. Cameron, G. shadle, J. blount & r. a. Dixon) signals for local and sys-temic responses of plants to pathogen attack. J. Exp. Bot. 55, 169–179.

(41) 2006 (With m.-b. Lascombe, n. buhot, b. bakan, D. marion, J.-p. blein & t. prangé) Crystallization of Dir1, a Ltp2-like resistance signalling protein from Arabidopsis thaliana. Acta Crystallogr. F 62, 702–704.

(42) 2008 (With J. Fan & C. Crooks) high-throughput quantitative luminescence assay of the growth in planta of Pseudomonas syringae chromosomally tagged with Photorhabdus luminescens luxCDABE. Plant J. 53, 393–399.

(43) (With m.-b. Lascombe, b. bakan, n. buhot, D. marion, J.-p. blein, V. Larue & t. prange) the struc-ture of ‘defective in induced resistance’ protein of Arabidopsis thaliana, Dir1, reveals a new type of lipid transfer protein. Prot. Sci. 17, 1522–1530.

(44) 2009 (With J. Fan, L. hill, C. Crooks & p. Doerner) abscisic acid has a key role in modulating diverse plant–pathogen interactions. Plant Physiol. 150, 1750–1761.



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Christopher John Lamb CBE. 19 March 1950 - Royal …rsbm.royalsocietypublishing.org/content/roybiogmem/56/189.full... · Christopher John Lamb Cbe 19 march 1950 — 21 august 2009