Before potatoes and rice, there were pulses

Although I spent most of my career working on potatoes and rice, my first interest was pulse crops or grain legumes. In fact the first pulse that I studied was the lentil (Lens culinaris Medik.) when I was an MSc student at the University of Birmingham from 1970-1971.

So why the interest in pulses?

It was surely the influence of one of my mentors, Dr Joe Smartt (right) at the University of Southampton where I was awarded my BSc in Environmental Botany and Geography in 1970. A geneticist who had studied groundnuts in Africa and at Southampton was working on Phaseolus beans, Joe taught a second year genetics course, and two in the third or final year, on plant breeding and plant speciation.

He published two seminal texts on pulses in 1976 and 1990.

It was Joe who ignited my interest in plant genetic resources, and encouraged me to apply for a place on the one year MSc course at Birmingham on Conservation and Utilization of Plant Genetic Resources (CUPGR). The course had been launched by the head of the Department of Botany, potato expert, and genetic resources pioneer, Professor Jack Hawkes, with the first intake of students commencing their studies in September 1969. I landed in Birmingham a year later.

My three year undergraduate course at Southampton was a stroll in the park compared to the intensity of that one year MSc course. We had eight months of lectures and practical classes, followed by written examinations at the end of May. Each student also had to complete a piece of independent but supervised research, and present a dissertation for examination in September. In order to take full advantage of the summer months, planning and some initial research began much earlier. First of all for most of us, we had to decide on a topic that was feasible and doable in the allotted time, and assemble the necessary seed samples ready for planting at the most appropriate date.

Almost immediately I decided on three points. First, I wanted to run a project with a taxonomy/natural variation theme. Second, I wanted—if feasible—to work on a pulse species. And finally (which I decided quite quickly after arriving in Birmingham) I wanted to work with Dr Trevor Williams (right) who delivered a brilliant series of lectures on variation in natural populations, among others.

Trevor and I thumbed our way through the Leguminosae (now Fabaceae) section of Flora Europaea, until we came upon the entry for Lens, and the topic for my project leapt off the page: Lens culinaris Medik. Lentil. Origin unknown.

My project had two components:

  • An analysis of variation in the then five species of lentil (one cultivated, the others wild species; the taxonomy has changed subsequently) from herbarium specimens borrowed from several herbaria in Europe. I also spent a week in the Herbarium at Kew Gardens in London taking measurements from their complete set of lentil specimens.
  • A study of variation in Lens culinaris from living plants, with seeds obtained from Russia (the Vavilov Institute in St Petersburg), from the (then) East German genebank in Gatersleben, and from the agricultural research institute in Madrid.

With the guidance of another member of the Botany department staff, Dr Herb Kordan, I made chromosome preparations and counts of all the Lens culinaris samples I’d obtained, confirming they were all diploid with 2n=2x=14 chromosomes. In the process, we developed a simple but effective technique for making chromosome squash preparations, and this led to my first ever publication in 1972. Just click on the title below (and others in this post) to read the full text.

In September 1971, I submitted my dissertation, Studies in the genus Lens Miller with special reference to Lens culinaris Medik. (which was examined by Professor Norman Simmonds who was the course External Examiner), and the degree was awarded.

I proposed that the wild progenitor of the cultivated lentil was Lens orientalis (Boiss.) Hand.-Mazz., a conclusion reached independently by Israeli botanist Daniel Zohary in a paper published the following year.

In 1971-1972, Carmen Kilner (née Sánchez) continued with the lentil studies at Birmingham, leading to a publication in SABRAO Journal in 1974. Our paper added further evidence to confirm the status of Lens orientalis.

When I began my lentil project, I had ideas to extend it to a PhD were the funding available. However, in February 1971 Jack Hawkes had just returned from a potato collecting mission to Bolivia, and told me about an exciting opportunity to spend a year in Peru at the newly-founded International Potato Center (CIP), from September that same year. My departure to Peru was delayed until January 1973, so I began a PhD on potatoes with Jack in the meantime. And with that move to potatoes, I assumed that any future work with pulses was more or less ruled out. However, from April 1981 I was appointed Lecturer in Plant Biology at Birmingham, and needed to develop a number of research areas. Would pulses figure in those plans?


While I wanted to continue projects on potatoes at Birmingham, I also decided to return partially to my first interest: pulses. And while I never had major grants in this area, I did supervise graduate students for MSc and PhD degrees who worked on a range of grain and forage legume/pulse species. Here I highlight the work of three students. There may have been more who worked on pulses, but after four decades I can’t remember those details.

Almost immediately after returning to Birmingham, I discovered (by looking through Flora Europaea once again) that the origin of the grasspea, Lathyrus sativus, was unknown. The grasspea is a distant relative of the ornamental sweetpea, Lathyrus odoratus, one of my favorite flowers since I was a small boy. My grandfather used to grow a multitude of sweetpeas in his cottage garden in Derbyshire. Anyway, I set about assembling a large collection of seed samples (or accessions) of grasspea and wild Lathyrus species from agricultural centers and botanic gardens worldwide.

The academic year September 1981-September 1982 was my first full year at Birmingham. Among the CUPGR intake was a Malaysian student, Abdul bin Ghani Yunus (right), who asked me to supervise his MSc research project. I persuaded him to tackle a study of variation in the grasspea and its wild relatives, much along the lines I had approached lentil a decade earlier.

We published this paper in 1984, and I guess it heralded what would become, a several decades later, an international collaborative effort to improve the grasspea and make it safer for human consumption.

Ghani returned to Malaysia, and I didn’t hear from him for several years. Then, in 1987, he contacted me to say he’d secured a Malaysian government grant to study for his PhD and would like to return to Birmingham. But to work on a tropical species, the name of which I cannot remember.

I persuaded him that would not really be feasible in Birmingham as we didn’t have the glasshouse space available, and it would be hit or miss whether we would be able to grow it successfully. I suggested it would be better to carry on his Lathyrus work from where he left off. And that’s what he did, successfully submitting his thesis in 1990 from which these papers were published.


Among the 1986 CUPGR intake was a student from Mexico, José Andrade-Aguilar (right) who was keen to attempt a pre-breeding study in Phaseolus beans, specifically trying to cross the tepary bean, Phaseolus acutifolius A. Gray with the common bean, Phaseolus vulgaris L.

José published two papers from his dissertation.

This next paper (for which I no longer have a copy) described how pollinations in Phaseolus species could be made more successful.


Then, in 1987, a student from Spain, Javier Francisco-Ortega (right, actually from Tenerife in the Canary Islands) joined the course, and he and I worked closely on his MSc and PhD projects until I left Birmingham to join IRRI in the Philippines in July 1991.

Javier was an extraordinary student: hard-working, focused, and very productive. After completing his PhD in 1992, he took two postdoctoral fellowships in the USA (at Ohio State University and the University of Texas at Austin) before joining the faculty of the Department of Biological Sciences at Florida International University in 1999, where he has been Professor in Plant Molecular Systematics since 2012.

For his 1988 MSc dissertation, Javier studied the variation in Lathyrus pratensis L., using multivariate analysis, and publishing this paper some years later.

Then, having successfully completed his MSc, and being awarded a second Spanish government scholarship, Javier began a PhD project to study the ecogeographical variation in an endemic forage legume from the Canary Islands, Chamaecytisus proliferus (L. fil.) Link., known locally as tagasaste or escobón, depending whether it is cultivated or a purely wild type.

With a special grant from the International Board for Plant Genetic Resources (IBPGR, now Bioversity International) in Rome, Javier returned to the Canary Islands in the summer of 1989 to survey populations and collect seeds from as many provenances as possible across all the islands, and I joined him there for several weeks.

Collecting escobón (Chamaecytisus proliferus) in Tenerife in 1989

After I left Birmingham, my colleague Professor Brian Ford-Lloyd took over supervision of Javier’s research, seeing it through to completion in 1992.

Together we published these papers from his research on tagasaste and escobón.

Once I was in the Philippines, I forgot completely about legume species, apart from contributing to any of the papers that were published after I’d left Birmingham.

One aspect that is particularly gratifying however is seeing the work Ghani Yunus and I did on Lathyrus still being cited in the literature as efforts are scaled up to improve grasspea lines.


 

Beets, ‘beans’, and Canaries

Lying off the Atlantic coast of northwest Africa by less than 600 miles, the Canary Islands archipelago comprises seven large islands, and a small group of islets off the north coast of Lanzarote, the island that lies furthest east and north. Volcanic in origin, and arid for the most part, their flora comprises many interesting endemic species found only on the Atlantic islands of MacaronesiaI’ve visited the Canaries twice, both in the 1980s, to collect plant germplasm (and also take a family holiday). Both expeditions were funded by the International Board for Plant Genetic Resources (IBPGR, now Bioversity International, based in Rome, Italy). So, as someone who studied potatoes and rice (and some legumes) most of my career, how did I become involved with collecting germplasm in the Canaries?

Brian Ford-Lloyd

Searching for beets
After leaving the International Potato Center in March 1981, I arrived at The University of Birmingham to begin my decade-long teaching career as Lecturer in Plant Biology from 1 April. Almost immediately, my colleague and fellow lecturer, Brian Ford-Lloyd (who retired a few years back as Emeritus Professor of Plant Conservation Genetics) invited me to join him on a collecting trip to the Canaries to look for wild relatives of beets (Beta spp.) that would contribute to an IPBGR global initiative on beet germplasm.

Now while I had my own experiences of germplasm collecting of cultivated (and some wild) potatoes in the Andes of South America between 1973 and 1976, I had no experience of beets whatsoever. Brian was keen to have me along on the trip because I did have one very important skill: I spoke (quite) fluent Spanish, and he expected that our Canarian counterparts would speak little English (which turned out to be more or less correct). So, not only would I be an experienced pair of germplasm hands, I could also be interpreter-in-chief.

Fortunately the dates for the trip coincided with my personal timetable then. Having arrived back in the UK at the end of March, my wife Steph (and daughter Hannah) stayed with her parents in Essex while I settled into my new job at the university, and while we house hunted. By the time Brian and I headed off to the Canaries in June, we’d bought our house, but moving in was not scheduled until the first or second weeks of July. So this was a great opportunity for me to join Brian.

Trevor Williams

Brian completed his PhD in 1973 under the supervision of Trevor Williams, submitting a thesis on the biosystematics of the genus Beta. As part of that research he made a collecting trip throughout Turkey in the early 1970s; and subsequently he maintained his research interest and activity in beets. Collecting in the Canaries was part of an IBPGR global initiative on beets.

Our particular interest there was a group of three beet species of Beta Sect. Patellares (I’m not sure if, or how, the taxonomy of Beta has changed in the intervening years) native to the archipelago, little represented at that time in different germplasm collections. Beets were reported from a range of localities throughout the islands, most often around the coasts or in ruderal habitats, but rarely inland (except in Fuerteventura) where the terrain is too high. In any case, this beet germplasm was considered under threat of genetic erosion, and had to be collected before habitats were lost through expansion of tourist resorts and holiday homes. Brian tells me he has been back to some of the sites where we collected and they have indeed been lost in this way.

Arnoldo Santos-Guerra

Travelling to the Canaries from Elmdon Airport (now Birmingham Airport) via London and Madrid, our first stop was Gran Canaria, staying for a couple of nights at the Jardín Botánico Canario Viera y Clavijo, where British botanist Dr David Bramwell was the director (and his wife Zoë, an acclaimed botanical artist). Those first days were essentially to find our feet, take some advice from David on where best to collect, before heading off to the island of Fuerteventura, the next island east from Gran Canaria, where we would meet our local expert and collaborator, Dr Arnoldo Santos-Guerra of the Centro Regional de Investigación y Tecnología Agrarias, Tenerife. For the collections in Tenerife, La Palma, and La Gomera we were joined by Arnoldo’s colleague, Lic. Manuel Fernández-Galván.

L-R: Brian, Arnoldo, Manuel, and me

In all, we collected 93 samples of beets from 52 locations on five islands: Gran Canaria, Fuerteventura, Tenerife, La Palma, and La Gomera.  Afterwards we published a trip report¹ in the FAO/IBPGR Plant Genetic Resources Newsletter.

On Tenerife, La Palma, and particularly La Gomera, there are precipitous inclines from the main roads down to the ocean’s edge. Deeply dissected landscapes ensure that wild beet populations are isolated from one another, even over relatively short distances as the cliff coastlines project into the ocean, with coves and beaches in between, where beets were often found. Therefore our ability to collect beet samples was quite often dependent entirely upon accessibility to the beach. The photos below were taken in Fuerteventura, Tenerife, and La Gomera. In some of them you can see the level of urbanization, almost 40 years ago, in many localities that were suitable environments for wild beets. The housing and tourist developments must be many times greater today.

But the actual process of collecting was not difficult at all, and seeds were often sampled from most if not all plants in some populations. Wild beets have a prostrate habit, and the ‘seeds’ were often found, in abundance, underneath the living plants. It was then just a question of scooping up handfuls of the seeds into a collecting bag, and annotating the collecting information appropriately.

Beta webbiana (left) and B. procumbens (right) from the Canary Islands

I say ‘seeds’, but the morphology of beets is a little more complex than that. Actually what we collected were small fruits with a hard pericarp, with several joined together to form multigerm seedballs. Modern sugar beet varieties are monogerm, a trait discovered in a wild beet species, in the former Soviet Union (Ukraine, in fact) during the 1930s . Because of their impermeability to moisture, and also due to the arid environments in which these beets species grew, we were confident that we were collecting viable seeds. In fact, as Brian explained to me, beet seeds are quite difficult to germinate.

Morphology of a beet inflorescence, seedballs, and a sugar beet (from: Wikipedia)

On our return to Birmingham, the seeds were added to the Birmingham Beta Collection that Brian curated, and other collections that are part of the World Beta Network. One recipient was Lothar Frese in Germany, now at the Julius Kühn-Institut in Quedlinburg. This germplasm has been used in a variety of studies looking at disease resistance such as Cercospora leaf spot resistance in B. procumbens in particular, and there has been much work since in terms of genetic mapping for resistance. After Brian retired, his beet collection was passed to the Genetic Resources Unit at the Warwick Crop Centre for safe storage.

A beet -‘bean’ linkage
In addition to beets, we collected 11 samples of other crops, among which was just one sample of a shrub or tree fodder legume, tagasaste, from La Palma, classified botanically as Chamaecytisus palmensis, and cultivated by many farmers. In our trip report, referred to above, we commented that the species did seem to be quite variable and, given its wider potential as a fodder legume, we suggested that it would warrant further study.

Javier Francisco-Ortega

And that was the last I thought about tagasaste until six years later when a young Spanish student from Tenerife, Javier Francisco-Ortega, enrolled on the genetic resources MSc course at Birmingham. Thirty years ago this month! I supervised Javier’s MSc dissertation on chromosome variation in Lathyrus pratensis, one of around 150 species in a genus that also contains the commonly-grown garden sweetpea, L. odoratus, and the edible grasspea L. sativus that was one of my research interests during the 1980s.

Anyway, to cut a long story short, Javier was an outstanding student, and began a PhD project with me in October 1988 on the ecogeography of the tagasaste complex, now classified taxonomically as C. proliferus. Only the forms from La Palma are popularly known as tagasaste (the ‘C. palmensis‘ we’d seen in La Palma in 1981), whereas those from the rest of the archipelago are commonly called escobón.

Morphological variants of tagasaste and escobón, Chamaecytisus proliferus

Tagasaste is the only form which is broadly cultivated in the Canary Islands and, since the late 19th century, also in New Zealand and Australia (particularly as fodder for sheep and goats). It has also become naturalized in Australia (South Australia, New South Wales, Victoria and Tasmania), Java, the Hawaiian Islands, California, Portugal, North Africa, Kenya, Tanzania and South Africa.

When I resigned from the university in June 1991 to join the International Rice Research Institute in the Philippines, supervision of Javier’s PhD passed to Brian.

In Spring and Summer 1989, and with funding from IBPGR, Javier began a systematic survey of 184 tagasaste and escobón populations throughout the archipelago (all islands except Fuerteventura and Lanzarote which are too dry), taking herbarium samples from each for morphological study, and revisited later to collect seeds. I joined Javier in July to assist with the collection of seeds from the Tenerife populations. Our trip report² was published in Plant Genetic Resources Newsletter in 1990. Arnoldo Santos-Guerra and Manuel Fernández-Galván were also contributors to this work.

Escobón populations are found commonly growing in gullies among pine forests, and appear to thrive here where there is the ever-present expectation (and danger) of forest fires. Indeed periodic burning appears to support the maintenance of escobón populations. These photos show the habitats of escobón populations in Tenerife, and Javier and myself making collections.

While more common in La Palma, farmers in Tenerife grow a few bushes of tagasaste in their terraces (seen on the right edge of the field in the picture below) on the north-facing slopes of the Teide volcano sloping down to the Atlantic.

We deposited duplicate seed samples in the Spanish national genebank in Madrid, and also in Tenerife. Javier took seeds back to Birmingham for further study, especially for analysis of molecular variation. Besides his PhD thesis, submitted successfully in 1992, his research led to several other scientific papers on morphological variation, phytogeography, ecogeographical characterization, genetic diversity, and the history of origin and distribution.

After he completed his PhD at Birmingham, Javier took postdoctoral fellowships at Ohio State University and the University of Texas at Austin before returning to Tenerife for a couple of years. In 1999 he was appointed Assistant Professor in the Department of Biological Sciences at Florida International University in Miami. He became Full Professor in 2012. He also has a joint appointment at the Fairchild Tropical Garden just south of Miami, as head of the Fairchild Plant Molecular Systematics Laboratory, with a special interest in cycads and palms, as well as an abiding interest in island floras. He has maintained his links with Arnoldo Santos-Guerra and David Bramwell.

In this video, Javier talks about his interests and the impact of his botanical research.

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¹ Ford-Lloyd, B.V., M.T. Jackson & A. Santos Guerra, 1982. Beet germplasm in the Canary Islands. Plant Genetic Resources Newsletter 50, 24-27.

² Francisco-Ortega, F.J., M.T. Jackson, A. Santos-Guerra & M. Fernández-Galván, 1990. Genetic resources of the fodder legumes tagasaste and escobón (Chamaecytisus proliferus (L. fil.) Link sensu lato) in the Canary Islands. Plant Genetic Resources Newsletter 81/82, 27-32.