Obituary: FRANK ROUND (1927–2010)

Hannelore Håkansson, the late Bjørg Stabell and Frank Round at the 14^th International Diatom Symposium in Japan, in 1996 (courtesy Dr M.B. Edlund; Norman Andresen appears behind on the left).

Two obituaries for Frank Round have already appeared in print, one by Dick Crawford (2011) and another by Eileen Cox (2011). These pay handsome tribute to his achievements and qualities and include some recollections by his students, colleagues and acquaintances. Further information has been given by Brian Moss in the foreword to a Festschrift published in 2006 (Crawford et al. 2006). It would be wrong, however, if we failed to commemorate Frank's life and work in the journal that he did so much to establish. This short article was therefore developed from a tribute to Frank Round presented at the 22nd International Diatom Symposium in Ghent, Belgium, 27–31 August 2012. I have tried to avoid repeating what Dick and Eileen have already written, but inevitably there is some duplication.

Frank Round died peacefully at his home on 25 October 2010 after a long illness. When I visited him a couple of months before he died, he was very frail following a fall, but we talked easily about the times before cheap transatlantic air travel, when he and his family travelled by ship to the USA for fieldwork and summer study visits. Frank travelled extensively and developed a wide network of contacts, huge experience and a wonderful reprint collection, which was mined by generations of research students. His publications often reflected his travels, e.g., studies of the phytoplankton of the Gulf of California, marine epipelon in Massachusetts, the benthic microalgae of Finnish lakes and pools, and the marine littoral diatoms of Hawaii. His interests were equally wide-ranging and he published significant papers on green algae and chrysophytes, as well as many on diatoms. His lectures to undergraduates spanned every taxonomic group and every field of algal biology, from physiology to phylogeny. In final-year phycology classes, we were set to work to prepare seminars on topics in oceanography, palaeoecology, cytology or biochemistry, on dinoflagellates, charophytes or red algae, and Frank seemed to be a master in all. This could have been intimidating if it had not been coupled with a kindly manner and personal anecdotes and insights (‘as I was driving to work this morning, I was thinking about\ldots ”), which made it clear that phycology was full of opportunity and interest, with plenty still to be discovered and understood.

Frank Round was born in the English Midlands in May 1927. Hence he was old enough to remember the post-Depression years of the 1930s but too young for military service in World War II. He grew up in Birmingham and went to university there, gaining a first class Bachelor's degree in Botany in 1948, after which he began a PhD on freshwater benthic microalgae. This was also based in Birmingham but with a period of fieldwork at the Freshwater Biological Association laboratories, which were then located in the mock-gothic Wray Castle, a mansion close to the western shore of Windermere in the English Lake District. His supervisor there was J.W.G. Lund, who was at that time engaged in his classic studies of freshwater phytoplankton (e.g., Lund 1949, 1954) but had earlier worked on benthic microalgae during his own PhD studies (Lund 1942). In turn, John Lund had himself been supervised by the pre-eminent British phycologist of the first half of the twentieth century, F.E. Fritsch, and the foundation for both John Lund's and Frank's first studies can perhaps be traced to Fritsch's masterly review of freshwater algal ecology in 1931. Frank's first university post was in the University of Liverpool but, after a short period back in Birmingham, he settled in the University of Bristol and remained there until and after his official retirement (early, so that he could get on with research!) in 1987.

Though many people probably remember Frank most for the systematic and taxonomic works he published in the second half of his life, Frank's PhD studies were ecological and he always regarded himself primarily as an ecologist. A full list of Frank's publications, assembled for his Festschrift (Crawford et al. 2006), shows that, among the 176 books, peer-reviewed papers, symposium papers and chapters he wrote, there are approximately equal numbers (c. 75 apiece) dealing with ecology and palaeoecology, on the one hand, and diatom morphology and taxonomy, on the other. Frank's first paper (Round 1953) examined the influence of sediment type and water column on the microalgae growing in the sediments of Malham Tarn in northern England. In it, he introduced a trapping method for semi-quantitative assessment of sediment algae, which he used for a number of ecological studies in the 1950s and was later refined for investigations of rhythmic migrations in the 1960s (Palmer & Round 1967). In the 1950s and 1960s, once based at the University of Bristol, Frank reported on microalgal communities from lake and river sediments, sand dunes, soils, bryophytes and elsewhere, in terrestrial, freshwater and marine environments. The significance of his work on lake epipelon is underlined by its prominence in volume 3 of the Treatise on Limnology (Hutchinson 1975). There were also palaeoecological studies of lake and oceanic sediments. However, although Frank's papers usually contained estimates of abundance and often dealt with seasonality, he never applied more than a bare minimum of statistical analysis and the increasing emphasis in ecology on ‘rigorous’ quantitative analysis was possibly one reason why Frank's research moved increasingly towards systematics from the 1970s onwards.

Unlike most British ecologists of the mid-20th century, Frank favoured the continental phytosociological approach, referring approvingly to the Braun–Blanquet school of ecologists. He took especial pains to clarify existing community terminology (e.g., epipelon, endopelon, epilithon) in several key articles (Round 1956, 1971a), and added a new term ‘epipsammon’ (Round 1965a) for the organisms living attached to sand grains, a community that particularly interested him. He repeatedly emphasized the need for proper classification of algal communities and wrote in The Ecology of Algae, ‘if one believes, as I do, in the concept of communities as collections of species living together and recurring in spatially separated habitats … then it is as essential to identify and describe these communities as it is to identify and describe the constituent species’ (Round 1981, p. 31). His books and papers include many tables listing and quantifying the species present in natural habitats. However, in case anyone should think that Frank was interested only in vegetational classification, they should note another sentence in the same section as the above quotation, in which he wrote that ‘the identification and description of the community is no more nor less important than its functional aspect’ (the italics are Frank's).

During the 1960s, a new tool became available to biologists: the scanning electron microscope (SEM). By good fortune, one of the first to make extensive use of the SEM was one of Frank's neighbours in the University of Bristol, the entomologist Howard Hinton FRS, who allowed Frank to use his Cambridge Stereoscan instrument. Figure 1–4 is a digitized image from the first numbered SEM negative in Frank's archive and foreshadows his early 1970s papers on aspects of the valve and girdle structure of Stephanodiscus and Cyclotella. I have not yet been able to find a date for this photograph, but it was probably taken in late 1967. The first SEM photograph that Frank published was one of Stephanodiscus used by Daniel Jackson for the cover and frontispiece images of a symposium volume (Jackson 1968) that contained chapters derived from a conference held in 1967. Characteristically for this period of his life, Frank's chapter was not on diatom morphology but on the effects of light and temperature on algae and included discussion of his contemporary work on vertical migration rhythms. The Jackson symposium Stephanodiscus image (Frank's copy of this is shown in Figs 2–4) may or may not have been the first published SEM photograph of a diatom, but even if it was not, Frank was certainly among the pioneers in this new field, together with Grethe Hasle (who published the first scientific papers on diatoms using the SEM in December 1968: Hasle 1968a, 1968b) and Bob Ross and Pat Sims at the Natural History Museum in London.

Figure 1—4. Archived scanning electron micrographs by F.E. Round (now in the Royal Botanic Garden Edinburgh). Fig. 1. Digitized image from negative #1 in Frank Round's collection, which was probably taken in late 1967 or early 1968. Fig. 2. A sheet of early SEM photographs of Stephanodiscus (printed photographs in Frank's collection are arranged by genus). Figs 3–4. Enlargements of Fig. 2, showing the micrograph (from negative 135) used as a frontispiece by Daniel Jackson (1968) and Frank's notes about the image, in his unmistakable handwriting (Fig. 4). The specimen came from Comber Mere in Cheshire, England (the information published by Jackson was incorrect).

Among Frank's early SEM papers, his 1970 paper on Hantzschia virgata is a good example of what he aimed to achieve with the new technology and deserves to be better known. The paper set Hantzschia in the context of other genera belonging to the same family, one of which (Cylindrotheca) had recently been revised by Reimann & Lewin (1964) using transmission electron microscopy of cultures; reported detailed light microscope (LM) observations of the frustule and chloroplast structure of a particular Hantzschia population at Barnstable, Massachusetts, sampled during one of Frank's many foreign trips; examined type material of H. virgata; gave a full account of the valves and girdle bands using the SEM; reconstructed the frustule in drawings to clarify the structure for readers; and finally, through reference to his observations of cell division in Hantzschia, addressed the distinction between Hantzschia and Nitzschia (and affirmed the likely validity of their separation), which had been questioned by Lauritis et al. (1967). Thus all aspects of the Hantzschia were examined and synthesized to produce a complete account of its morphology and taxonomy. But to understand Frank, it is important to realize that this work grew out of a quite different study – an examination of the vertical migrations of H. virgata in the sediments of Barnstable, Massachusetts, in which Palmer & Round (1967) had demonstrated a remarkable endogenous tidal rhythm, which still intrigues students and whose mechanism is still unknown.

The SEM was seductive, however, and during the 1970s and 1980s, Frank spent increasingly less time on ecology and increasingly more on morphology and taxonomy, and this led to his most highly cited contribution, The Diatoms (Round et al. 1990: currently with over 1550 citations in Web of Science). Frank had the idea of producing an ‘atlas’ of diatoms at around the time that I left Bristol (1978) and I remember his confidence even then that he, Dick Crawford and I already had almost enough material to produce a comprehensive account of the genera. He had underestimated the task and it was not until 10 years later that, with Gill Lockett's secretarial help in drafting and redrafting the text, we completed the plates, drawings and typescript and sent them all to Cambridge University Press. In the intervening years, the atlas had mutated into a more ambitious work, which used SEM and cytological data to change the diatom classification rather than merely illustrate existing taxa, and it had gained a long introduction to explain diatom structure and biology. All through the long gestation of The Diatoms, Frank drove the project forward, writing letters, getting new material and photographs when gaps in our coverage became evident (though of course, many remained unfilled in the final volume), producing the first drafts of many generic descriptions and introductory chapters, so that we had something to work on, and reassuring Cambridge University Press that we would complete the task.

As a senior academic in the University of Bristol, Frank sat on a number of Boards and Committees and did not avoid his share of administration. However, rather than discussing what ought or might be done to improve this or that, Frank preferred to get on with doing. He worked to establish the International Society for Diatom Research, founded Diatom Research and edited it for many years, and also produced many excellent volumes of Progress in Phycological Research, establishing his own publishing house (Biopress) with his wife Marie and Gill and Tony Lockett, when he felt that the existing publisher of Progress was pricing it so high that no one would buy it. Among other Biopress publications was the beautiful Freshwater Algae by John Lund and his wife Hilda (Canter-Lund & Lund 1995), which Frank was able to publish at a manageable cost despite the wealth of superb colour photographs that it contains. Throughout his long working life, Frank kept in close touch with John Lund, who has outlived him and is due to celebrate his 100th birthday in 2012.

It was characteristic of Frank that he did not add his name to papers by his PhD students, unless he had made a major contribution himself. For example, all the credit for the classic work on diatom locomotion by the late Lesley Edgar (Edgar & Pickett-Heaps 1984) and the earlier work on the same topic by Margaret Harper (Harper & Harper 1967) remained with them. Frank made no joint publications at all with either of these, or with Eileen Cox, Penelope Dawson, Andrew Leitch, Paul Broady and Bulent Sen (PhDs on tube-dwelling diatoms, diatom cytology, charophytes, terrestrial Antarctic algae, and plankton parasites, respectively). Dick Crawford and I never published with him on our PhD topics (Melosira sensu lato and the Bacillariaceae, respectively). Today, such a policy might be suicidal (since research leaders need to prove their credentials in order to retain staff and funding), but it was not so in the 1960s and 1970s, yet even then Frank's generosity was unusual. Frank was the sole author for over 100 of his papers and books.

Though he was sometimes critical in private of some researchers and had strong opinions about the worthlessness of some contemporary biology, his comments during symposia were generally positive and encouraging, and he was prepared to spend a lot of time helping to improve manuscripts, especially for those whose first language was not English. A conversation with him was usually a tonic. In retrospect, he must surely have got tired of cheering up other people; it did not seem so at the time. I rarely saw Frank angry but he had little patience with anyone he thought was wasting their opportunities, or who seemed to be trying to create an élite. His drive to get things done could create resistance and sometimes animosity, but on his part I detected no rancour and he bore no grudges. In his publications, the characteristics of his academic father and grandfather (John Lund and F.E. Fritsch) can be detected: a huge breadth of knowledge and a belief in scholarship (valuing and evaluating the contributions of others, old or new), openness to new ideas (though to the regret of many of us, he never became computer-literate), an international perspective, insights from personal experience (valuable and otherwise unpublished observations are to be found even in his student textbook, The Biology of Algae, 1965b), and the ability to synthesize and generate new ideas. Frank was not afraid to ask for advice, as can be seen from the acknowledgements in his papers, which often record that the manuscript had been read and criticized by others, and he was not worried about being proved wrong: to be corrected was all part of making progress. And sometimes he was indeed quite wrong, as in his early attempts to understand girdle structure in Stephanodiscus (Round 1971b).

Frank was sociable, liked good food and wine, and was a keen gardener. He was married to Marie, who unfortunately became ill and died only a year or so after Frank; they are survived by their three daughters, Nicola, Kathryn and Clare. Many phycologists got to know Marie and the children either during visits to their home, ‘The Orchard’, where visitors received a generous welcome, or when the whole family went on some of Frank's earlier working ‘holidays’ abroad.

Finally, two images to wrap up this piece and leave with you. One will have to be imagined because, as far as I know, no photograph was taken: Frank in the rain, dressed in suit and tie (he dressed quite formally at the university) and raincoat, wading into Priddy Pond in his wellington boots to obtain sediment and epipelon for a student practical with a ‘Round tube’ (a c. 1.5-m glass tube, c. 12 mm internal diameter, introduced to phycology by Round 1953). The second image is the one that begins this article, which was very kindly supplied by Mark Edlund and shows Hannelore Håkansson, the late Bjørg Stabell and Frank enjoying life in Japan in 1996, during the 14th International Diatom Symposium. Frank was inspiring and encouraging and fun. He leaves good memories.

Note on collections

Frank Round's slide preparations were transferred to the Royal Botanic Garden Edinburgh when Frank cleared his room in the University of Bristol in the late 2000s. They contain most of the material used in his publications, together with an extensive collection made during Frank's late work on UK rivers, which was reflected in unusually few publications. Unfortunately, almost all the unmounted material could not be rescued and had to be thrown away. Frank's LM and SEM negatives, and his reprint collection, are also currently housed in the Royal Botanic Garden.