"In the United States, the University of Georgia has been preeminent
in its contribution to our knowledge of the Ascomycetes, with
Professors Julian Miller and E. S. Luttrell developing a system of
classification that emphasizes the structure of the ascus and the
ascocarp centrum ..."
The above quote, taken from the textbook INTRODUCTORY MYCOLOGY by Alexopoulos and Mims (1979), reflects the reputation that the University of Georgia has acquired during the past nine decades for the research conducted on the taxonomy and morphology of ascomycetes and other fungi by investigators at the University. This article will trace the development of this research and its impact on taxonomic mycology.
The first mycologist employed in the Georgia Agricultural Experiment Stations was Dr. Bascombe B. Higgins. He joined the staff of The Georgia Experiment Station, located near Griffin, on October 1, 1913, as Botanist and Head of the Botany Department, a position that marked the beginning of research in mycology and plant pathology in the state. Dr. Higgins had recently completed his Ph.D. degree in mycology under Dr. George F. Atkinson at Cornell University, where he acquired a life-long interest in the fungi. As the first and only plant pathologist in the state, Dr. Higgins was confronted by many pathological problems which required his attention, but he still found time to devote to his special interest, the developmental morphology of ascomycetes. Between 1913 and 1936 Dr. Higgins conducted morphological studies on a number of ascomycetes of importance to plant pathology. His earliest studies were on the genus Coccomyces, in which he described C. hiemalis and C. lutescens as the sexual states of Cylindrosporium diseases of stone fruits, and also conducted studies on ascocarp development in both species. He then turned his attention to the genus Mycosphaerella, many species of which cause serious leafspot diseases of a variety of economically important plants. Among the species described and studied developmentally were M. bolleana on fig, M. personata on grape, M. tulipiferae on Liriodendron (tulip poplar), and M. persica on peach. These studies still stand as some of the most detailed conducted on these genera and they often appear in mycology and plant pathology textbooks as examples of such studies. Another important series of studies were those on the physiology and parasitism of Sclerotium rolfsii, the cause of white mold disease of peanuts and many other crops, in which he obtained the basic information that was later utilized in developing cultural control methods for this severe pathogen. In later years Dr. Higgins turned his attention to peanut breeding and other areas of research. Colleagues recognized his mycological accomplishments by naming Colletotrichum higginsianum, Taphrina higginsii, Piricularia higginsii, and the genus Higginsia, in his honor. His contributions to agriculture were acknowledged with the naming of 'Higgins', a new cultivar of bronze muscadine grape.
Mycological research was continued at Georgia Experiment Station by Dr. Wilbert A. Jenkins, who joined the staff on July 1, 1934. He was particularly interested in leafspot fungi and described a number of new ascomycetes as causal agents of some of these important diseases, including Mycosphaerella angulata, cause of angular leafspot of muscadine grape. During his seven years at the Station he also produced some excellent morphological studies. Especially noteworthy were his descriptions of the important leafspot pathogens of peanut, Mycosphaerella arachidicola and M. berkeleyii, cause of the early and late leaf blights, respectively, and the developmental studies he conducted on these species. His study of centrum development in Cordyceps agariciformia still ranks as one of the best on this genus.
At the main campus of the University of Georgia in Athens, Dr. Julian H. Miller returned to his position in the Department of Botany in 1928 after receiving the Ph.D. degree in mycology from Cornell University. Five years later he became the Chairman and only faculty member of the Department of Plant Pathology in the College of Agriculture.
Dr. Miller's mycological interests were very broad, and during the next three decades, he made numerous contributions to mycological knowledge. He collected extensively in the state and established a herbarium for these and additional specimens received from other collectors or by exchanges with other herbaria. At the time of his retirement nearly 10,000 specimens were housed in the herbarium. The Georgia specimens formed the basis for a series of papers on Georgia pyrenomycetes, in which several new species were described.
Of more fundamental importance, however, were his studies on the life histories and morphology of various ascomycetes. His papers on Myriangium, Bagnisiopsis, and Phyllachora are of special interest in this regard, for it was from these studies that he concluded that developmental morphology and the internal structure of the ascocarp provided a far more fundamental basis for classifying these fungi than did earlier criteria. In 1949 he published a classification scheme for the ascomycetes embodying many of these concepts, and this was the first work to employ developmental data in the separation of the major groups of the pyrenomycetes, a large subgroup of ascomycetes. Of particular significance was his demonstration that two separate lines of development occurred among the perithecial ascomycetes, which corresponded to the Ascohymeniales and Ascoloculares types of ascocarp development originally proposed by the Swedish mycologist J. A. Nannfeldt in 1932. This resulted in rearranging a number of genera of perithecial ascomycetes.
Throughout his career Dr. Miller maintained an especial interest in members of the Xylariaceae. His numerous publications on these fungi established him as a world authority on the family. From these studies came a world monograph of the genus Hypoxylon, which he completed after his retirement in 1958. He died before it could be published and it was brought out posthumously. At the time of his death in 1961 he was also working on a monograph of Xylaria, but unfortunately it was not sufficiently advanced to permit completion by others.
In addition to his many publications and professional reputation, Dr. Miller left as a legacy the large collection of fungi, mostly ascomycetes, that he assembled during his career. This collection has been designated the Julian H. Miller Mycological Herbarium (GAM); it now contains approximately 25,000 specimens, among them several types and many of the specimens he examined during his monographic studies. The Miller Herbarium was originally housed in the Department of Plant Pathology of The University of Georgia, but in the mid-1980s it was transferred to the newly formed Museum of Natural History. Dr. Miller's contributions to the University were recognized by the naming in 1976 of the new Plant Sciences Building (depicted below) in his honor. The genus Jumillera also has been named in his honor.
Associated with Dr. Miller during most of his career was Dr. George E. Thompson. Also a graduate of Cornell, he joined the Department of Plant Pathology at Athens in the fall of 1937. During their years together the two men collaborated in the collection and study of Georgia pyrenomycetes, and co-authored several papers describing them, including several new species. He also was an ardent collector and assembled a sizable collection of fungi that is now part of the Miller Herbarium.
Dr. Thompson's own special area of interest was in fungi that cause tree diseases, and he devoted many years of study to this subject. He described several new tree disease fungi from Georgia, with emphasis on those causing leaf diseases. Especially noteworthy were his studies on diseases of poplar, during which he described the life cycles of several species, including Neofabraea, Pleuroceras, and Guignardia, all pathogenic on poplar. In later years Dr. Thompson made important contributions to our knowledge of the rust fungi in Georgia, and he was actively studying them at the time of his death in November, 1960.
A year after the resignation of Dr. Wilbert A. Jenkins from the Georgia Experiment Station, Dr. Everett S. Luttrell joined the staff on October 1, 1942, as Associate Botanist. Shortly thereafter he began publication of a series of papers on the developmental morphology of ascomycetes that soon established him as a leading authority in this field. Dr. Luttrell had received the Ph.D. degree from Duke University, where he studied under Dr. Frederick A. Wolf. His early studies included the genera Sphaerostilbe (Nectria), Phacidium, Morenoella, Myiocopron, Stomiopeltis, Ellisiodothis, and Dothidea.
In 1947 Dr. Luttrell moved to the University of Missouri as Assistant Professor of Botany. He soon found that the heavy teaching load left insufficient time for research and in 1949 he returned to the Georgia Experiment Station. It was while at Missouri, however, that he completed work on the book for which he would become widely known.
The publication in 1951 of Taxonomy of the Pyrenomycetes represented a major development in the classification of the ascocarpic ascomycetes. Following a review of the literature on the morphology of the perithecial ascomycetes Dr. Luttrell concluded that there are a limited number of ways in which ascocarps develop, and that the perithecial ascomycetes could be separated into natural groups on the basis of the method of development and the structure of the centrum. Following Miller and Nannfeldt, he recognized two separate lines of development for the perithecial ascomycetes, which he termed the Euascomycetes and the Loculoascomycetes. These classes corresponded to the Ascohymeniales and Ascoloculares developmental types of Nannfeldt. He also made an important correlation of these two types of development with the manner of ascus dehisence; species with Ascohymeniales development have unitunicate asci, and species with Ascoloculares development have bitunicate asci. The impact of this new approach can be seen in a comparison of the classification schemes for these fungi used by Alexopoulos in the first (1952) and second (1962) editions of INTRODUCTORY MYCOLOGY (see table below). One result of using these new criteria was the transfer of certain well-known pathogens to different subclasses and the rearrangement of numerous genera.
------------------------------------------------------------------------- First Edition (1952) Second Edition (1962) EUASCOMYCETES EUASCOMYCETIDAE Plectomycetes Plectomycetes Aspergillales Eurotiales *Myriangiales #Microascales Pyrenomycetes Pyrenomycetes Erysiphales Erysiphales Sphaeriales Meliolales Hypocreales Chaetomiales Laboulbeniales Sphaeriales *Microthyriales Hypocreales *Dothideales Clavicipitales *Pseudosphaeriales Diaporthales *Hysteriales Coryneliales Coronophorales LOCULOASCOMYCETIDAE *Myriangiales *Pleosporales *Dothideales *Microthyriales *Hysteriales * = Orders containing species with bitunicate asci. # = Included in the Sphaeriales in first edition. -------------------------------------------------------------------------
The basic classification scheme proposed by Dr. Luttrell for the perithecial ascomycetes was predominant for three decades, during which it was recognized by the majority of mycologists and followed in most major textbooks on mycology.
The recent application of molecular data to ascomycete taxonomy indicates that some of Luttrell's concepts are no longer valid, but these data do support the existence of two separate developmental patterns for the ascomycetes, as proposed by Nannfelt, Miller, Luttrell, and others. Information on the development and structure of ascocarps, however, continues to be important in understanding the biology of these fungi.
Following publication of his book, Dr. Luttrell continued developmental studies of ascomycetes, but he also initiated studies on the taxonomy of graminicolous species of Helminthosporium, a large genus of conidial fungi with sexual states belonging to the ascomycetes. By applying the principles of developmental studies to conidium ontogeny in the Helminthosporium complex, he demonstrated that small and seemingly insignificant differences in conidium structure and conidiophore behavior supported the division of this genus into four separate genera. He also demonstrated that these differences could be correlated with the sexual states, several of which he discovered and described as new species. This led eventually to his publication of a classification scheme for the conidial fungi in which the major taxonomic separations are based largely on conidium ontogeny. In later years he became interested in the manner in which plant parasitic fungi developed within the host, conducting studies on ergot in grains and of several smut fungi and their hosts. These studies combined the techniques of mycology and plant pathology, thus bridging the gap between these two areas of investigation.
In September, 1960, Dr. Richard T. Hanlin was employed as Assistant Plant Pathologist at the Georgia Experiment Station to collaborate with Dr. Luttrell on developmental studies in ascomycetes. Dr. Hanlin had recently completed his degree in mycology under Dr. Lewis E. Wehmeyer at the University of Michigan. Although initially employed in a temporary position, he was later chosen to fill the position left vacant by the retirement of Dr. Higgins. While at the Georgia Station, Dr. Hanlin published developmental studies on species of Nectria, Hypocrea, and Hypomyces, which clarified the pattern of ascocarp ontogeny in the order Hypocreales. Following the emergence of the mycotoxin problem in food crops in the early 1960s, he became a member of the regional research project to determine the prevalence of aflatoxigenic fungi in peanuts, and later, in pecans, both important crops in Georgia. During the next six years several thousand isolates from the southeastern United States belonging to the Aspergillus flavus group were examined and identified, along with numerous other fungi. These studies greatly increased our knowledge of the fungal biota in peanuts and pecans, two major crops in Georgia. In addition, to strengthen the mycology curriculum at the main campus in Athens, he commuted twice weekly one quarter per year to teach an advanced mycology course, since there was no mycologist in Athens at the time.
In July, 1965, Dr. Luttrell left the Georgia Experiment Station to become head of the statewide Department of Plant Pathology at the University of Georgia. A year later Dr. Hanlin transferred to the Athens campus, thus ending the mycology research program at Griffin. This was a time of expansion in the sciences at the University of Georgia, as the department was enlarged and a doctoral program was added. Dr. Hanlin revived the introductory mycology course originally taught by Dr. Miller, and later added courses on the biology of the ascomycetes and the identification of fungi. During the next 30 years he and his students conducted developmental studies on a number of ascomycetes, including Leptosphaerulina, Sporormiella, Melanospora, Sordaria, Gelasinospora, Podospora, Cercophora, Didymella, and Phyllachora. In the 1980s he began working on tropical plant pathogenic ascomycetes in collaboration with researchers in Brazil and Venezuela, and later in Mexico. During this time he also developed aids for the identification of these fungi and later, in collaboration with former student Miguel Ulloa of the National University of Mexico, they published an Illustrated Dictionary of Mycology in both English and Spanish. In 1962 Dr. Hanlin was designated as curator of the mycology herbarium established by Dr. Miller. In 1978 the herbarium became a unit of the Museum of Natural History and in 1986 the holdings were physically transferred to the Natural History Museum. Following his official retirement from the Department of Plant Pathology in June, 2001, Dr. Hanlin has continued to serve as volunteer curator of the mycology herbarium, which is now housed in the Museum Annex, a part of the Administrative Services Warehouse. His work has been recognized with the naming of the genus Hanliniomyces and the species Capronia hanliniana and Dendrostilbella hanlinii.
With the expansion of the faculty, Dr. Kenneth E. Papa was employed in 1964 to develop a program in the genetics of filamentous fungi. Dr. Papa received his degree in Neurospora genetics under Dr. Adrian M. Srb at Cornell University. He initially continued his research on Neurospora at Georgia, but soon turned his attention to mycotoxigenic fungi. He demonstrated the presence of a parasexual cycle in Aspergillus flavus and A. parasiticus, and studied the genetic makeup of these fungi. He was actively engaged in these studies at the time of his death in 1986.
The genetics position was filled by Dr. David D. Pope, who developed a research program on the genetics of the corn smut fungus, Ustilago maydis. In 1993, however, he left this position to become a computer programer with industry.
In 1995 Dr. Scott E. Gold was employed for the fungal genetics position. Since that time he has developed an active research program on the molecular genetics of Ustilago maydis.
In 1986 Dr. William E. Timberlake was employed in a joint position in the Departments of Genetics and Plant Pathology to study the genetics of filamentous fungi. Dr. Timberlake specialized in the molecular genetics of Aspergillus nidulans, and he and his students made numerous contributions to this area. In 1993 he left Georgia to work in industry.
In 1986 E. S. Luttrell retired from the University of Georgia after nearly 44 years of service. In the later years of his career he received many honors, both from professional societies and from the University. His mycological colleagues honored him with the naming of the genus Luttrellia, the species Biolaris luttrellii, Physalacria luttrellii and Pringsheimia luttrellii, and the dedication of a symposium volume, Ascomycete Systematics - The Luttrellian Concept. He was honored by the University with a Distinguished Professorship and by the E. S. Luttrell Lecture Series that was established in the Department of Plant Pathology.
The vacancy created by Dr. Luttrell's retirement was filled by Dr. Charles W. Mims. Dr. Mims received his training at the University of Texas under Dr. Constantine J. Alexopoulos, and he came to Georgia recognized as an outstanding teacher and electron microscopist. At Georgia he applied ultrastructural techniques to the study of host-parasite interactions and spore development in plant pathogenic fungi. These studies added a new dimension to the mycology program and its interface with the study of plant pathogens by others in the Department.The first person trained in mycology employed in the Department of Botany at the University of Georgia was John Moore Reade, a native of Canada, who was hired as Instructor in Botany in 1907. Two years later, upon completion of his doctoral degree at Cornell University, he was promoted to Professor. His dissertation was on the taxonomy of the genus Sclerotinia, the species of which are important plant pathogens of commercial crops. Among the new species he described was Sclerotinia vaccinii-corymbosi (now placed in the genus Monilinia), an important pathogen of blueberries in Georgia and elsewhere. Although Dr. Reade made some collections of fungi after returning to Georgia from Cornell, he soon turned his attention to the taxonomy of phanerogams and did not publish more in mycology. He did make an important contribution to mycology, however, by serving as major advisor to Julian H. Miller during his research for the M.S. degree.
The next mycologist employed in the Department of Botany was Dr. Lindsey S. Olive, who was hired as Assistant Professor in 1945. Dr. Olive had received his doctoral degree from the University of North Carolina three years earlier under Dr. William C. Coker. Dr. Olive studied the jelly fungi of Georgia, but left after only one year because the heavy teaching load precluded doing research. This was unfortunate for Georgia, as Dr. Olive went on to become one of the preeminent mycologists in the United States during a long and distinguished career.
Twenty-two years elapsed before the Department of Botany employed another mycologist. Dr. Melvin S. Fuller was hired as head of that department in 1968. Dr. Fuller received his degree from the University of California, Berkeley under Dr. Ralph W. Emerson. Dr. Fuller's research interests were in the Chytridiomycetes, especially the mode of spore development and the ultrastructure of their zoospores. He and his students were successful in cultivating and studying a number of these minute aquatic fungi and elucidating their biology. Dr. Fuller retired from the University of Georgia in 1995.
A year after Dr. Fuller came to Georgia, he employed a second mycologist in the Department of Botany with the addition of Dr. David Porter in 1969. Dr. Porter received his degree from the University of Washington under Dr. Howard Whisler. Dr. Porter's interests are in the ecology and biology of zoosporic marine fungi and fungus-like organisms, especially species of Labyrinthula that parasitize seagrasses. He also has worked with endolithic fungi that live in marine shells.
Dr. Alan Jaworski came to Georgia as a post-doctoral fellow in plant physiology in 1969, after receiving his doctorate under Dr. Albert Siegel at the University of Arizona. By the end of his post-doctoral stay he had become interested in fungal physiology and was hired in 1971, when he initiated research on Blastocladiella emersonii, an aquatic fungus. He continued these studies until his death in January, 2000.
In 1974 Dr. William E. Barstow became the third mycologist in Botany. Dr. Barstow received his doctorate from Purdue University under Dr. James Lovett. Dr. Barstow's research involves the use of light and electron microscopy to study the life cycles and development of nonfilamentous water molds. In recent years both Drs. Jaworski and Barstow have been heavily involved in teaching elementary biology to undergraduates.
The position left open by Dr. fuller's retirement was filled by Dr. Michelle Momany, who came to Georgia in 1996. Dr. Momany received her doctorate in Microbiology from the University of Texas. In her research she applies the techniques of molecular biology to the study of the development and organization of the cytoskeleton and cell wall in Aspergillus nidulans and A. fumigatus.
The addition of the mycologists in the Department of Botany greatly broadened the mycology program at the University of Georgia by providing specialists and research in groups and areas not previously represented.
Professional recognition of the quality of mycological research by University of Georgia faculty members is demonstrated by the election of five of them (J. H. Miller, E. S. Luttrell, M. S. Fuller, R. T. Hanlin and C. W. Mims) to the presidency of The Mycological Society of America.
In addition to the mycologists mentioned above, various other individuals in the University and related federal agencies also work with fungi as part of their research. Thus the University of Georgia continues to be a center for mycological research in the United States.
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