125 years of laboratory research in botany at The New York Botanical Garden DENNIS WM. STEVENSON New York Botanical Garden, 2900 Southern Blvd., Bronx, NY 10458-5126, USA; e-mail:
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Abstract. A brief history of laboratory research in botany is given with a focus on trends through time, impact on botanical research at The New York Botanical Garden, and its impact on the teaching of botany in university settings. Key Words: Cullman Program for Molecular Systematics, genomics, botanical history, paleobotany, plant anatomy, plant physiology
Laboratory approaches to research in botanical science have been a part of the science program at The New York Botanical Garden since the institution’s founding in 1891. At the time, laboratory research programs at botanical gardens were unusual, both in the United States and abroad, with the notable exceptions of the Royal Botanic Gardens, Kew, and botanical gardens in Germany, particularly in Berlin and Munich. In the discussion that follows, I examine the cause of this unique situation in the founding of The New York Botanical Garden, the leadership role the institution has played in North America, and its impact on botanical research and education. There were three factors that influenced the decision to develop a botanical research laboratory at The New York Botanical Garden. The first is the result of the intersection of the Garden’s foundation in a university setting; second, the creation of NYBG was a New World example of the latest trends in Europe; and third, very importantly, is due to the foresight of two of the founders, Nathaniel Lord Britton and Henry Hurd Rusby, who wanted to create a world class botanical garden with a broad range of programs—from public to scientific. The need for a research botanical garden was deemed a necessity by N. L. Britton, then Chair of the Botany Department at Columbia University, as well as by other faculty studying plants, particularly H. H. Rusby in the College of Pharmacy. In addition, I believe that the role of Columbia University in the then history of North American Botany was a factor particularly through the work of John Torrey in the Flora of North America and John Strong Newberry in Paleobotany.
It was the confluence of these many factors that led to the inclusion of a laboratory program and facilities at NYBG. The Royal Botanic Gardens, Kew was the model for the organization of The New York Botanical Garden in 1891 (Gleason, 1996), and at that time Kew had established the Jodrell Laboratory where research in plant anatomy, paleobotany, and plant chemistry were being pursued. The research scientists of The New York Botanical Garden also served as faculty members in botany at Columbia University, which housed the herbarium, library, paleobotanical collections, and laboratories until the Museum Building at NYBG was completed in 1899. Another factor was N.L. Britton’s desire to bring the latest thinking, expertise, and knowledge of the ongoing advances going on in German botany to the students and faculty at Columbia University. Thus, he hired Emily Lovira Gregory in 1889 (E. Britton, 1897) as the first woman professor in the history of Columbia University, albeit at Barnard College. Britton did this because Emily Gregory had done her doctoral work in Germany with the outstanding plant anatomists and physiologists of the day, and thus was well trained in the modern aspects of these areas of botany and represented a fundamental shift in botanical science thinking (Gregory, 1887, 1892; Schmid & Stevenson, 1987; Rudolph, 1996). Botany courses were originally taught at Columbia from 1891 to 1899 only subsequently moved to the new Museum Building at NYBG (Lentz & Bellengi, 1996). From that point, Columbia’s laboratory classes were also moved to the Garden, necessitating laboratory facilities at NYBG (N. Britton, 1915; Fig. 1A, B). Additionally, Emily Gregory (1895)
Brittonia, DOI 10.1007/s12228-016-9418-4 ISSN: 0007-196X (print) ISSN: 1938-436X (electronic) © 2016, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
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FIG. 1. Original lab (A, B) and current Plant Science Research Laboratory (C, D). A, B. Original lab on the 6th floor of the Museum building (current location of the LuEsther T. Mertz Library), with close-up of mycology research area (B). C, D. Plant Science Research Laboratory, built in 2006.
had published her BElements of Plant Anatomy,^ the first textbook published on the subject in North America (Schmid, 1987; Rudolph, 1984); Rusby (1895) had published a general botany textbook; and Rusby and Jelliffe (1895) had published a text book emphasizing plant chemistry and physiology. These all became contemporary textbooks, some of which went through many editions (Rusby & Jelliffe, 1895, 1899; Rusby, 1911, 1924). Emily Gregory died prematurely in 1897 and was succeeded by her master’s degree student, Carlton Curtis, who completed a doctorate at Syracuse University and then Cambridge before returning to the faculty at Columbia. He took up where Gregory left off and published leading textbooks (Curtis, 1897, 1907, 1931), one of which went through many editions. These placed NYBG as a leader in that period’s modern integrated botany, a trend that has been continuous and still exists today. This point is developed in the following discussion via the examination of the careers of the prominent scientists involved in laboratory research during their careers at NYBG. Daniel Trembly MacDougal, the first director of the laboratories, came to NYBG in 1899 from the University of Minnesota. He was basically a plant physiologist who pioneered the field of ecological plant physiology, now known as ecophysiology. He
was brought to NYBG in keeping with the institution’s goal of maintaining the forefront of the new botany, particularly for having published a textbook on experimental plant physiology (MacDougal, 1895). He was the leading American authority on desert ecology and one of the earliest botanists to research chlorophyll. He was the inventor of the MacDougal dendrograph, used to record changes in the volume of tree trunks. MacDougal became Director of Botanical Research at the Carnegie Institution in Washington, DC in 1907. In 1950, he was elected honorary president of the International Botanical Congress in Stockholm and was awarded the first Certificate of Distinguished Service from the NYBG in 1956. Alexander P. Anderson spent only a few years at NYBG. He was a master’s student under MacDougal at the University of Minnesota and then completed his doctorate in Munich. He arrived at NYBG in 1899, and in 1901, he discovered that starch granules could be heated within a tube and blasted into powder with the resulting particles smaller than starch granules themselves. Later, rice, wheat, and other whole grain cereals were treated in the same way, producing the puffed cereals that we still consume today. Over 1000 experiments were conducted on different products in 1902. He left NYBG later that year,
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after patenting the process, and later started the Anderson Puffed Rice Company in 1905. Anderson developed a partnership with Quaker Oats, and Quaker Oats took over production of the puffed cereals. Paleobotany played an important role in laboratory studies involving collections. The New York Botanical Garden’s collection of plant fossils is one of the founding collections for North American paleobotany, resulting from the work of John Strong Newberry, who became Chair of Geology and Paleontology at Columbia College in 1866. Newberry had a strong influence on N. L. Britton and Arthur Hollick and thus influenced the formation of NYBG. It was during Newberry’s term that the Columbia Geology Museum was established. After Newberry’s death in 1892, Columbia’s fossil plant collection was transferred to NYBG. Significant collections of late Cretaceous floras from New England, New Jersey, and New York were added to it when Columbia Professor Arthur Hollick became a curator at NYBG. Hollick also collected many Tertiary fossil plants from Puerto Rico and Cuba, integrating paleobotanical data with information on the extant flora that was being studied and published at the same time. After 55 years of a paleobotanical hiatus, Herman Becker in 1958 succeeded Hollick as curator in paleobotany and spent his career studying the Tertiary floras of Montana until his retirement in 1974. In the 1970s, Karl Niklas was the last paleobotanist at NYBG. He continued the tradition of remaining on the forefront by utilizing phytochemical methodology to determine some of the chemical components of fossilized plants when they had been living. In the absence of a paleobotanist on staff, The New York Botanical Garden’s paleobotanical collection was transferred in its entirety to the Division of Paleobotany at Yale’s Peabody Museum in 1983. Without paleobotanical collections, NYBG’s preeminence in paleobotany ceased. Marshal Avery Howe was one of the original faculty who received his doctorate from Columbia in 1898 and then joined the Columbia faculty and NYBG. He moved to NYBG in 1901, when the herbarium was moved to the Garden. He worked mainly with marine red algae, including their life cycles in culture, and the morphology and structure of liverworts. When he passed away in 1936, after 35 years at NYBG, research on algae ceased for the next 70 years until the modern era with the arrival of Kenneth G. Karol and his molecular approaches and culturing of the Charales.
Another early component of laboratory activities was mycology, for which a detailed history is available for the first 100 years (Rogerson & Samuels, 1996), so I provide only highlights here. William Alphonso Murrill was the first full-time mycologist who joined the staff in 1904. Murrill published numerous important monographs and over 500 scientific articles on a wide range of botanical subjects. Most importantly, he identified the pathogenic fungus, Diaporthe [Cryphonectria] parasitica, on American Chestnut trees, which causes Chestnut Blight disease, the worst ecological disaster that mankind has ever inflicted on North America. The presence of the Chestnut Blight in North America was the direct result of human activities via the importation of diseased saplings from Asia to North America. Murrill’s contemporary at NYBG was Fred Jay Seaver, who came to NYBG in 1908 as Director of Laboratories when MacDougal went to the Carnegie Institution. Another contemporary was Bernard Ogilvie Dodge, who received his doctorate from Columbia University in 1912 and then served at the Bureau of Plant Industry of the U.S. Department of Agriculture in Washington, DC as a plant pathologist until coming to NYBG in 1928. B.O. Dodge was very much what Rogerson and Samuels (1996) termed an experimental mycologist. He was quite interested in genetics and life cycles, and he worked out the details of the life cycle of Neurospora. He demonstrated regularities in the Mendelian inheritance that made Neurospora a critical taxon for geneticists. Dodge’s findings on Neurospora and his association with Edward Tatum led to the latter receiving the Nobel Prize in 1958. B.O. Dodge was the only person Tatum thanked for developing the basic research organism that allowed Tatum to understand the role of genes in biochemical processes, which in turn led to his Nobel prize. Dodge also published a plant pathology textbook with the Garden’s bibliographer, H. W. Rickett (Dodge & Rickett, 1943), which went through two editions, plus a third edition with Pascal Pirone and then another two with Pirone alone (1978). Pascal Pompey Pirone joined the staff of the NYBG in 1947, succeeding B. O. Dodge as resident plant pathologist. Pirone had come to NYBG from Rutgers University, where he had published several books on horticulture and plant pathology, the latter of which were coauthored with B. O. Dodge and H. W. Ricketts. Pirone’s
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horticulture books on tree maintenance (1948, 1952, 1978) were also widely used as textbooks, one of which went through six editions (1988). Again the laboratory programs were providing textbooks in the tradition of modern research for science and education. NYBG honored Pirone with its Distinguished Service Award in 1982. Another contemporary of B. O. Dodge was Arlow Burdette Stout. Stout and Dodge were graduate students together at Columbia, receiving doctorates a year apart (1913 and 1912, respectively). Stout was appointed Director of the Laboratories in 1911, a position he held until his retirement in 1947. Stout was a geneticist who worked on the genetics of intraspecific self- and cross-incompatibilities in the sexual reproduction of flowering plants with emphasis on seedless grapes, poplar trees, and daylilies. He produced many hybrid cultivars of all three groups and published over 350 papers in his 36 years at NYBG. In 1954, he was awarded the NYBG Distinguished Service Award. The years that Dodge and Stout spent as graduate students and Columbia faculty must have been extremely exciting. In addition to these two working at NYBG and teaching at Columbia, Columbia also had on its faculty several pioneers in genetics, including Thomas Hunt Morgan (a Nobel Prize winner), Leslie Dunn, and Theodosius Dobzhansky in Zoology, and Edmund Sinnott and Robert Harper in Botany. Genetics was at the forefront of research at NYBG during this very important period of its history. As we shall see, presently, after a 60 year absence, genetics is again a very important part of the Garden’s Science program. In 1938, William Jacob Robbins became the Director-in-Chief of NYBG, equivalent to the role of President in contemporary times. Robbins was a renowned plant physiologist who came to the Garden from the University of Missouri where he was the Dean of the Graduate School, former head of the Botany Department, and Acting President of the University. Robbins was also a member of the National Academy of Sciences, and while Director-in-Chief at NYBG, he was also the Chair of the Botany Department at Columbia University. He was the first director who specialized in plant physiology and microbiology rather than taxonomy. There was apparently much consternation expressed by the staff on Robbins’ appointment as Director-in-Chief because he was known as a plant physiologist and not a taxonomist. I particularly like Henry Allan Gleason’s (1947) response to those critics:
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BThere’s been no real change. The Twentieth Century goes to Chicago every night, no matter who the conductor is.^ Robbins’ primary research interests were tissue culture, normal and teratological growth, the role of plant hormones in plant development, and the physiology of aging in higher plants, as well as the nutritional requirements of filamentous fungi and their secondary chemicals. He was particularly interested in research on an experimental program to extract antibiotic substances from basidiomycetes, discussed in Rogerson and Samuels (1996). Robbins also had a great interest in plant genetics and was a close colleague with Stout and Dodge. Robbins retired in 1958 just after a new laboratory building opened in 1956, which later was named the Charles B. Harding Laboratory in commemoration of Harding's thirty years of service to the Garden on its Board of Directors, and Robbins having refused to have it named after himself (Kavanagh & Hervey, 1991). Robbins, in keeping with NYBG tradition, produced university textbooks on botany (Robbins & Rickett, 1939). After Robbins retired, Richard Klein joined the NYBG faculty in 1958 with a research program in physiology of crown gall disease and secondary plant compounds. This began an association with Arthur Cronquist, which resulted in the establishment of a program in phytochemistry and the publication of an interesting book on the taxonomic and evolutionary significance of plant chemistry (Klein & Cronquist, 1967). Cronquist was very supportive and knowledgeable of laboratory approaches to botany. His relationship with his colleagues allowed him to produce text books in his own right (Cronquist, 1961, 1973) for university botany courses that went through multiple editions and that were translated into Spanish. Because of Cronquist’s influence, phytochemistry became an important component of the laboratory research with the hiring of David Giannasi as a phytochemist in the early 1970s. At this same time, Karl Niklas arrived as paleobotanist. Together, Giannasi and Niklas applied phytochemisty to paleobotany with surprising and groundbreaking success. During the late 1970s and early 1980s, complicated financial situations led to the loss of this type of research program at NYBG, but it was reinstituted in the late 1980s with new faculty, P. Mick Richardson and Fred Seaman, carrying out phytochemical research until 1990. Meanwhile, Bassett Maguire and associates were actively pursuing plant anatomy as a source of data
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FIG. 2. The Plant Science Research Laboratory houses the Lewis B. and Dorothy Cullman Program for Molecular Systematics, Plant Genomics, Structural Botany, Phytochemistry, and Mycology.
for the comparative anatomy of the Clusiaceae as well as the Rapataceae and Xyridaceae. This work continued for many years throughout the 1970s and well into the 1990s. The resulting microscope slides produced by this program are part of the laboratory’s permanent collections (see Campbell, 2016, this issue). Tetsuo Koyama was in charge of the anatomical program of NYBG’s laboratories through the 1970s to the late 1980s pursuing anatomical studies on grasses and sedges in the main. The Harding Laboratory was closed for two years, 1984–1986 for asbestos abatement. Laboratory research ceased during that period, and the laboratory staff consisted of only P. Mick Richardson. In 1987, Ghillean Prance brought me from Barnard College of Columbia University to NYBG to reinstitute laboratory research. I had made my laboratory at Barnard College available to students and staff of NYBG for the period of closure, and I quickly reinstituted structural botany and phytochemisty at the Garden. I published a textbook in plant anatomy (Cutler et al., 2008), harkening back to Emily Gregory over one hundred years ago. The use of secondary plant compounds in both plant systematics and in biodiversity prospecting intersected so that funding was available and obtained both from the National Science Foundation and from major pharmaceutical companies. Thus, the results from the systematics helped determine what plants to investigate pharmaceutically and vice versa. This was an era of expansion of laboratory approaches to economic botany and plant systematics. I had also been pursuing molecular approaches to plant systematics at Barnard College. In
keeping with NYBG’s history of always pursuing and incorporating new approaches, in 1994, through the extraordinary generosity of Lewis and Dorothy Cullman, the Lewis B. and Dorothy Cullman Program for Molecular Systematics was created and plant genetics was then brought back via the Program for Plant Genomics. Molecular data was at that time coming into the forefront in systematics across biology and the Cullman Program represented collaboration between NYBG and the American Museum of Natural History. NYBG was a pioneer in using molecular approaches to investigating the evolutionary history of plants and fungi. The early years were spent using restriction site approaches and then later gene sequencing and currently whole genome sequencing. Research using molecular approaches to plant systematics and building and understanding the Tree of Life has been and is still pursued successfully across the land plants, freshwater algae, and fungi. Moreover, NYBG stayed at the forefront of bioinformatics expertise with the research of Damon Little, who developed DNA Barcoding methodologies for many applications, from identification of plant materials from archeological remains to the composition of dietary supplements. The facilities of the Cullman program are now used by all of the taxonomists at NYBG under the direction of Gregory Plunkett. In 2000, an initiative in Plant Genomics was begun at NYBG. In 2001, NYBG formed the New York Plant Genomics Consortium with Cold Spring Harbor Laboratory, and New York University and, in 2004, the American Museum of Natural
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History was added. The addition of the Plant Genomics program brought back plant genetics to laboratory research at NYBG and simultaneously added the evolution of development (evo/devo) to structural botany through the hiring of plant geneticist Barbara Ambrose. The data produced by the Cullman Program for Molecular Systematics is used to build evolutionary histories of plant groups, and those histories in turn are used to identify features of interest in plant evolutionary biology, agriculture, etc. This sets the stage for evo/devo to discover the genes responsible for the traits of interest and how those genes can be used in plant biology to explain how the features arose and diversified. During the late 1990s, it became clear that with the advent of these new approaches, the Harding Laboratory had become obsolete and a new facility would eventually be needed. As a result, in 2006, a new Plant Science Research Laboratory building was built to house Structural Botany, the Cullman Program, Plant Genomics, Phytochemistry, and Mycology (Figs. 1C, D, 2). This history has yet to be written, but suffice it to say that NYBG remains on the forefront of utilizing and integrating new approaches in laboratory research in botanical science.
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———. 1892. The Two Schools of Plant Physiology as at Present Existing in Germany and England. The American Naturalist 26: 211–217, 279–286. ———. 1895. Elements of plant anatomy. Ginn & Co., Boston, MA. Kavanagh, F. & A. Hervey. 1991. "William Jacob Robbins 1890-1978". Biographical Memoirs 60: pp. 292–328. National Academy of Sciences, Washington D.C. Klein, R & A. Cronquist. 1967. A consideration of the evolutionary and taxonomic significance of some biochemical, micromorphological, and physiological characters in the thallophytes. Quarterly Review of Biology, Volume 42. Lentz, D. L. & M. Bellengi. 1996. A brief history of the Graduate Studies Program at The New York Botanical Garden. Brittonia 48: 404–412. MacDougal, D. T. 1895. Experimental plant physiology. Henry Holt & Co., New York. Pirone, P. P. 1948. Maintenance of shade and ornamental trees. 2nd Ed. Oxford University Press, New York. ———. 1952. Complete guide to modern gardening. Hart Pub. Co., New York. ———. 1978. Diseases and pests of ornamental plants. 5th Ed. John Wiley, New York. [Note that the 3 Edition was with Dodge & Rickett and with Pirone as senior author]. ———. 1988. Tree maintenance. 6th Ed. Oxford University Press, New York. Robbins, W. J. & H. W. Rickett. 1939. Botany: a textbook for college and university students. 3rd Ed. Van Norstand, New York. Rogerson, C. T. & G. J. Samuels. 1996. Mycology at The New York Botanical Garedn, 1895-1996. Brittonia 48: 389–398. Rudolph, E. D. 1984. The first American plant anatomy book and its author. American Journal of Botany 71(5, pt. 2): 107–108. ———. 1996. History of the Botanical Teaching Laboratory in the United States. American Journal of Botany 83: 661– 671. Rusby, H. H. 1895. Botany. D.O. Haynes, New York ———. 1911. A manual of structural botany: an introductory text-book for students of science and pharmacy. Lea & Feibinge, Philadelphia, PA. ———. 1924. A manual of structural botany: an introductory text-book for students of science and pharmacy. 2nd Ed. Revised. D.O. Haynes, New York ——— & S. E. Jelliffe. 1895. Essentials of vegetable pharmacognosy. A treatise on structural botany. Part I. The gross structure of plants. By Henry H. Rusby. Part II. The minute structure of plants. By Smith Ely Jelliffe. D.O. Haynes, New York ——— & ———. 1899. Morphology and histology of plants: designed especially as a guide to plant-analysis and classification, and as an introduction to pharmacognosy and vegetable physiology: Part I. The morphology of plants / by Henry H. Rusby. Part II. Plant histology / by Smith Ely Jelliffe. D.O. Haynes, New York Schmid, R. 1987. Annotated bibliography of works by and about Emily Lovira Gregory (1841–1897). Bulletin of the Torrey Botanical Club 114: 319–324. ——— & D. W. Stevenson. 1987. "Botanical text books," an unpublished manuscript (1897) by Emily Lovira Gregory (1841–1897) on plant anatomy textbooks. Bulletin of the Torrey Botanical Club 114: 307–318.