Annual Review of Phytopathology - Volume 34, 1996

Helen Hart was a Professor of Plant Pathology at the University of Minnesota from 1924 until retirement in 1966. Born in Janeville, Wisconsin, she died at Grants Pass, Oregon. Her scholarly research concentrated on wheat stem rust to understand host pathogen relationships and to develop rust–resistant cultivars. She did not teach formal courses but was heavily involved in making seminars a vital part of instruction, in teaching languages needed for graduate studies, and as an informal advisor for most rust research theses. She had common sense, excellent scientific judgment, and sound instincts on personnel matters that served the department well. A talented science writer, Hart served as editor of hundreds of theses and departmental manuscripts for publication. Her writing and editing skills were used as associate editor of Phytopathology for two years and as editor-in-chief from 1944–1951. A strong advocate of The American Phytopathological Sociecty, Helen Hart served on Council for 12 years and as President in 1956. Helen Hart was a great professional scientist who had a far-reaching impact on plant pathology during the twentieth century.

Swiss-born Dr. Gotthold Steiner was a pioneer in formulating the discipline of nematology. He worked with the American nematologist NA Cobb and together they were responsible for acceptance of the concept of nematode phytoparasites. Steiner had special expertise in anatomy, morphology, phytonematology, marine nematodes, nutrition, mermithids, and selected invertebrate taxa. He authored 191 scientific papers, established the ubiquitous phytoparasitic genus Helicotylenchus, described the pinewood nematode, and did significant work with three important economic pests, Ditylenchus dipsaci, Heterodera rostochiensis, and H. schachtii. He was responsible for introducing training programs in nematology in USDA laboratories.

The Red Queen Hypothesis (RQH) explains how pathogens may maintain sexual reproduction in hosts. It assumes that parasites become specialized on common host genotypes, reducing their fitness. Such frequency-dependent selection favors sexual reproduction in host populations. Necessary conditions are that resistance and virulence are genotype specific so that host genotype frequencies respond to changes in pathogen genotype frequencies, and vice versa. Empirical evidence on the genetic basis of disease, variation in resistance and virulence, and patterns of infection in sexual and asexual plants support certain features of the hypothesis. However, gene-for-gene interactions are generally not consistent with the RQH because they do not result in cycling of gene frequencies, unlike a matching allele mechanism. A conclusion of whether the RQH can explain the maintenance of sexual reproduction cannot be reached at present. Nevertheless, the RQH theory has shed light on many aspects of plant/pathogen interactions important for reducing pathogen damage in agricultural systems.

The scarcity of literature in the realm of plant clinics operating in the developing world brought us to the decision to perform a survey in non-OECD countries in Africa, Asia and Oceania, Latin America and the Caribbean, and in central and southern Europe. Forty-one completed questionnaires that refer to 79 plant clinics were returned. The best-equipped clinics are operated by the international research bodies. By and large, this review shows that considerable progress has been achieved in the physical development of plant clinics in developing countries and in staffing them with well-trained personnel. Not surprisingly, however, many inadequacies and inconsistencies still need to be addressed before clinics can fully play their part in providing plant protection services to their clients. Success and constraining indicators in the diagnostic performance of the clinics are identified. A model of the‘ideal plant clinic’ is suggested.

Dwarf bunt is a disease of wheat caused by the smut fungus Tilletia controversa Kuhn. Winter wheat (Triticum aestivum L.) is the primary host of economic significance. Although the total acreage affected by dwarf bunt is small relative to total wheat production worldwide, the disease has assumed attention disproportionate to its economic impact because it has become a matter of contention in world trade in cereals. This review describes the political and economic issues underlying the study and identification of T. controversa.

Thirty soilborne viruses or virus-like agents are transmitted by five species of fungal vectors. Ten polyhedral viruses, of which nine are in the family Tombusviridae, are acquired in the in vitro manner and do not occur within the resting spores of their vectors, Olpidium brassicae and O. bornovanus. Fungal vectors for other viruses in the family should be sought even though tombusviruses are reputed to be soil transmitted without a vector. Eighteen rod-shaped viruses belonging to the furo- and bymovirus groups and to an unclassified group are acquired in the in vivo manner and survive within the resting spores of their vector, O. brassicae, Polymyxa graminis, P. betae, and Spongospora subterranea. The viral coat protein has an essential role in in vitro transmission. With in vivo transmission a site in the coat protein-read through protein (CP-RT) of beet necrotic yellow vein furovirus determines vector transmissibility as does a site in a similar 98-kDa polyprotein of barley mild mosaic bymovirus. The mechanisms by which virions move (or are moved) into and out of the protoplasm of zoospores or of thalli needs study.

Epichloë species and their asexual descendants (Acremonium endophytes) are fungal symbionts of C3 grasses that span the symbiotic continuum from antagonism to mutualism depending on the relative importance, respectively, of horizontal transmission of sexual spores versus vertical clonal transmission in healthy grass seeds. At least seven sexual Epichloë species are identifiable by mating tests, and many asexual genotypes are interspecific hybrids. Benefits conferred by the symbionts on host plants include protection from biotic factors and abiotic stresses such as drought. Four classes of beneficial alkaloids are associated with the symbionts: ergot alkaloids, indolediterpenes (lolitrems), peramine, and saturated aminopyrrolizidines (lolines). These alkaloids protect host plants from insect and vertebrate herbivores, including livestock. Genetic engineering of the fungal symbionts as more suitable biological protectants for forage grasses requires identification of fungal genes for alkaloid biosynthesis, and DNA-mediated transformation of the fungi.

Numerous bacteria have been isolated from within plants, and many reported from xylem, but only three species of xylem-limited bacteria (XLB) that are fastidious in cultural requirements, are plant pathogens, and exclusively occupy xylem, have been well characterized. Two XLB, Xylella fastidiosa and Pseudomonas syzygii, are transmitted by sucking insects that feed on xylem sap but are not transmitted mechanically from plant to plant. In contrast, Clavibacter xyli is mechanically transmitted to plants by cutting tools. All of these XLB occupy a highly specialized yet diverse ecological niche: the water-conducting systems of an extremely wide range of plant hosts. A variety of detection methods are available as diagnostic aids; each method has advantages and disadvantages; no single method is best for all uses. Molecular and genetic comparisons of strains of XLB lag behind progress being made for many other plant-pathogenic bacteria, but such studies are needed to answer important questions: (a) How do XLB move from cell to cell within plants? (b) What are the physiological and genetic bases of plant host specificity for XLB? (c) Why are only xylem-feeding specialists vectors of X. fastidiosa (and probably P. syzygii), when many leafhoppers feed regularly (but not continuously) on xylem?

Although more than 30 bacterial avirulence genes have been cloned and characterized, the function of the gene products in the elictitation of resistance is unknown in all cases but one. The product of avrD from Pseudomonas syringae pv. glycinea likely functions indirectly to elicit resistance in soybean, that is, evidence suggests the gene product is an enzyme involved in elicitor production. In most if not all cases, bacterial avirulence gene function is dependent on interactions with the hypersensitive response and pathogenicity (hrp) genes. Many hrp genes are similar to genes involved in delivery of pathogenicity factors in mammalian bacterial pathogens. Thus, analogies between mammalian and plant pathogens may provide needed clues to elucidate how virulence gene products control induction of resistance.

The role and functioning of the anterior chemosensory organs of plant parasitic nematodes is examined, with particular emphasis on the amphids. The morphology of the amphids is discussed primarily in the context of the changes in the ultrastructure associated with different life stages. The involvement of amphidial secretions in chemoreception and the behavioral and electrophysiological analyses of nematode responses to semiochemicals are discussed with special reference to research on sex pheromones. These research techniques, combined with the use of lectins and antibodies, provide information on nematode sensilla that may lead to novel control strategies for economically important plant parasitic nematodes based on perturbing nematode sensory perception to prevent host or mate location.

In small-scale, subsistence agriculture in the tropics and the subtropics, traditional farming practices have evolved that provide a sustainable means of reducing the incidence and damage caused by pests including nematodes. Other newer, cultural and low-input practices can also be successfully introduced in small-scale farming. In sustainable and subsistence farming systems, nematodes and some other pests can be managed by integrating different farming practices into four strategies: preventing the introduction and spread of nematodes; using direct, nonchemical, cultural and physical control methods; encouraging naturally occurring biological control agents; and maintaining or enhancing the biodiversity inherent in multiple cropping and multiple cultivar traditional farming systems to increase the available resistance or tolerance to nematodes.

A variety of noncirculatively transmitted viruses have evolved a vector transmission strategy that involves, in addition to virions, virus-encoded proteins that are not constituents of virions. These “helpers” and the genes encoding them have been characterized for viruses in the genera Potyvirus and Caulimovirus. Several lines of evidence support the hypothesis that these helpers act by mediating retention of virions in regions of the vector's alimentary tract from which they subsequently can be egested to initiate an infection. The possible advantage this convergently evolved strategy could confer to noncirculatively transmitted virus quasispecies is discussed.

The 15 known viruses that occur in rice are rice black-streaked dwarf, rice bunchy stunt, rice dwarf, rice gall dwarf, rice giallume, rice grassy stunt, rice hoja blanca, rice necrosis mosaic, rice ragged stunt, rice stripe necrosis, rice stripe, rice transitory yellowing, rice tungro bacilliform, rice tungro spherical, and rice yellow mottle viruses. This paper describes their geographical distribution, relation to vectors, infection cycles, field dispersal, and development, and lists recorded outbreaks of the viruses. Many rice viruses have become serious problems since rice cultivation has been intensified. Double-cropping of rice using improved, photo-insensitive cultivars of short growth duration has significantly influenced the incidence of these viruses.

Rice tungro, the most important virus disease of rice in South and Southeast Asia, is caused by a complex of two viruses, rice tungro bacilliform virus (RTBV) and rice tungro spherical virus (RTSV). RTBV is a plant pararetrovirus with bacilliform particles, the structure of which is based on T = 3 icosahedral symmetry cut across the threefold axis.The particles encapsidate a circular double-stranded DNA of 8 kbp that encodes four proteins. The current information on the properties, functions, and expression of these proteins is discussed, as is the evidence for replication by reverse transcription. Two major strains of RTBV have been recognized, one from the Indian subcontinent and the other from Southeast Asia. RTSV particles contain a single-stranded RNA genome of 12 kb that encodes a large polyprotein and possibly one or two smaller proteins. The properties and processing of the polyprotein are described and the resemblance to picornaviruses noted.

The development of plant virus gene vectors for expression of foreign genes in plants provides attractive biotechnological tools to complement conventional breeding and transgenic methodology. The benefits of virus-based transient RNA and DNA replicons versus transgenic gene expression include rapid and convenient engineering coupled with flexibility for expeditious application in various plant species. These characteristics are especially advantageous when very high levels of gene expression are desired within a short time, although instability of the foreign gene in the viral genome can present some problems. The strategies that have been tested for foreign gene expression in various virus-based vectors include gene replacement, gene insertion, epitope presentation, use of virus controlled gene expression cassettes, and complementation. Recent reports of the utililization of virus vectors for foreign gene expression in fundamental research and biotechnology applications are discussed.

New evidence confirms earlier postulates that root signals to shoots, including abscisic acid, nitrate flux, and cytokinins, modify whole plant growth processes including leaf expansion, stomatal behavior, and biosynthesis of photosynthetic enzymes. Root signals are thought to reflect soil water, nutrient, and mechanical attributes, as sensed by roots. Meristematic activities in root tips initiate changes in root architecture, modifying the soil volume subject to root uptake, and may provide multiple sensory and signaling capabilities. Knowledge of root signals regulating whole plant growth processes suggests new analytical and experimental tools for integrated analysis of plant phasic development, optimal growth, and ecological fitness.

Phytotoxic effects of ozone are described with emphasis on secondary plant metabolism. Numerous ozone-induced genes, enzymes and stress metabolites of antioxidative and phytopathological defense reactions have been discovered for herbaceous plants and forest tree species. Ozone induces reactions normally elicited by viral and microbial pathogens. The molecular basis (receptors, signal chains) for induction by ozone remains to be elucidated. The induced stress reactions seem to change plant predisposition to either enhanced tolerance or susceptibility for a second stressor. The following topics are discussed: ozone and biotic disease, the role of ozone on field sites and ozone limit values.

Infection structures of phytopathogenic fungi are modified hyphae specialized for the invasion of plant tissues. Initial events are adhesion to the cuticle and directed growth of the germ tube on the plant surface. At the site of penetration, appressoria are often formed that may have melanized walls and develop high turgor pressure to support the penetration process. The penetration hypha accumulates components of the cytoskeleton in the tip and secretes a variety of cell wall–degrading enzymes in a highly regulated fashion in order to penetrate the cuticle and the plant cell wall. This article reviews recent papers on the cytology, physiology, and molecular biology of the penetration process.