This booklet is designed as a quick guide for identifying wheat and triticale diseases in the field. It is intended primarily for agricultural researchers, technicians and farmers in developing nations, but will also be of value to others. The text comprises a brief description of the major wheat and triticale diseases, insect pests, nematodes, physiologic and genetic disorders, and mineral and environmental stresses. Complementing this text and as an aid to the identification are numerous color photographs 1/, drawings 2/ and a brief diagnostic key. In the case of specific causal organisms, the perfect stage name (when known) is followed by the imperfect stage name in parentheses. While most of the diseases, pests, disorders, or stresses included can be economically significant, some are not and are presented only because they are unique or might be confused with more economically significant problems.
-------------------------------------------------------------------------------- Symptoms Possible Disorder -------------------------------------------------------------------------------- Leaf or leaf sheath ------------------- *Raised pustules with yellow, Leaf Rust (Brown Rust) orange or black spores Stem Rust (Black Rust) Stripe Rust (Yellow Rust) Flag Smut *Superficial white, pinkish or Powdery Mildew grayish fungal tissue Scab (Head Blight) *Superficial dark fungal tissue Spot Blotch Black Point Black Molds *Dark fungal fruiting bodies Septoria Tritici Blotch within brownish grey lesions *Flecks Leaf Rust (Brown Rust) Stem Rust (Black Rust) Stripe Rust (Yellow Rust) Powdery Mildew Spot Blotch Barley Yellow Dwarf Thrips Mites Genetic Flecking *Spots Tan Spot Physiological Leaf Spot Melanism and Brown Necrosis Genetic Flecking Herbicide Damage *Lesions or blotches resembling Leaf Rust (Brown Rust) halos with constrasting colored Stem Rust (Black Rust) borders Stripe Rust (Yellow Rust) Tan Spot Physiological Leaf Spot Minor Element Deficiencies *Other types of blotches Fusarium Leaf Blotch (Snow Mold) Basal Glume Rot/Bacterial Leaf Blight Barley Yellow Dwarf Herbicide Damage Frost Damage *Yellow to grey-white exudate Black Chaff/Bacterial Stripe Basal Glume Rot/Bacterial Leaf Blight *Yellowing, chlorosis, necrosis, Flag Smut stunted growth Take-All Eyespot (Strawbreaker) Sharp Eyespot and Rhizoctonia Root Rot Common Root Rot, Foot Rot, and Crown Rot Sclerotium Wilt (Southern Blight) Barley Yellow Dwarf Armyworms, Cutworms, and Stalk Borers Wireworms Seed Gall Nematode Cereal Cyst Nematode Root Knot Nematode Nitrogen, Phosphorus, and Potassium Deficiencies Aluminum Toxicity *Reddening, stunted growth Barley Yellow Dwarf Nitrogen,Phosphorus, and Potassium Deficiencies *Deformed, curled, rolled, Flag Smut twisted or thickened leaves Downy Mildew and/or leaf sheaths Barley Yellow Dwarf Mites Seed Gall Nematode Minor Element Deficiencies Herbicide Damage *Chewed, tattered, eaten or Armyworms, Cutworms, and Stalk Borers missing leaves Slugs, Snails, Grasshoppers, and Crickets *Chewed longitudinal stripes Cereal Leaf Beetle Slugs, Snails, Grasshoppers, and Crickets Stem or leaf sheath ------------------- *Raised pustules with yellow, Leaf Rust (Brown Rust) orange, brown, or black spores Stem Rust (Black Rust) Stripe Rust (Yellow Rust) *Superficial white, pinkish, or Powdery Mildew grayish fungal tissue *Black, shiny fungal tissue Take-All beneath leaf sheath *Brownish, elongated distinct Eyespot (Strawbreaker) eye-shaped lesions Sharp Eyespot/Rhizoctonia Root Rot *Uniform tan-brown Common Root Rot, Foot Rot, and Crown Rot discoloration *Brown to black blotches or Septoria Tritici Blotch stripes Black Chaff/Bacterial Stripe Physiological Leaf Spot *Lodging, broken stems Spot Blotch Eyespot (Strawbreaker) Sharp Eyespot and Rhizoctonia Root Rot Hessian Fly Wheat Stem Maggot Sawfly Frost Damage Root or crown ------------- *Darkish rots, lesions Take-All Common Root Rot, Foot Rot, and Crown Rot *White to black fruiting bodies Sclerotium Wilt (Southern Blight) *White mycelium Common Root Rot, Foot Rot, and Crown Rot Sclerotium Wilt (Southern Blight) *Shiny black crown node Take-All *Deformed, knotted or stubby Cereal Cyst Nematode roots Root Knot Nematode Aluminum Toxicity *Attached white or brown galls, Armyworms, Cutworms, and Stalk Borers cysts or nodules White Grubs Wireworms *Severed or chewed roots Armyworms, Cutworms, and Stalk Borers White Grubs Wireworms Head ---- *Entire spike black, transformed Loose Smut into a powdery mass *Raised pustules with yellow, Leaf Rust (Brown Rust) orange, brown or black spores Stem Rust (Black Rust) Stripe Rust (Yellow Rust) *Superficial white, pinkish or Powdery Mildew greyish fungal tissue Scab (Head Blight) *Superficial dark fungal tissue Black Molds (Sooty Molds) *Brown, black or purple blotches Septoria Tritici Blotch or streaks on glumes Spot Blotch Tan Spot (Yellow Leaf Spot or Blotch) Black Chaff/Bacterial Stripe Melanism and Brown Necrosis *Yellow exudate Black Chaff/Bacterial Stripe Bacterial Spike Blight *Entire head white, forced Leaf Rust (Brown Rust) ripening, seeds shrivelled or Black Molds (Sooty Molds) absent Barley Yellow Dwarf Stink Bugs Wheat Stem Maggot Sawfly Minor Element Deficiencies Frost Damage *Head is twisted or otherwise Downy Mildew distorted, may not emerge from Bacterial Spike Blight leaf sheath Barley Yellow Dwarf Seed Gall Nematode Minor Element Deficiencies Herbicide Damage Frost Damage Seed ---- *Discolored with a change in shape, size and/or texture - Color black; seed transformed Loose Smut into a powdery mass - Color gray to black; seed Common and Dwarf Bunt (Stinking Smut) easily crushable and has Karnal Bunt (Partial Bunt) strong odor - Color brown to black; seed Ergot hard (p. 34) - Partially discolored; seed Karnal Bunt (Partial Bunt) contains black, powdery spores, strong odor when crushed *Seed discolored only Alternaria Leaf Blight Scab (Head Blight) Black Point Black Molds (Sooty Molds) Seed Gall Nematode Entire Plant ------------ *Patches of stunted or damaged Take-All plants throughout the field Eyespot (Strawbreaker) Sharp Eyespot and Rhizoctonia Root Rot Common Root Rot, Foot Rot, and Crown Rot Sclerotium Wilt (Southern Blight) Barley Yellow Dwarf White Grubs Cereal Cyst Nematode Salt Stress Moisture Stress Heat Stress *Dwarf clumping Hybrid Necrosis (Dwarf Clumping) --------------------------------------------------------------------------------
Fungi differ from other plants in that they have no chlorophyll and thus lack photosynthetic capability. Instead of manufacturing their own food, fungi absorb nutrients from either living or dead host tissue. Fungi are dispersed in many ways: they may be seed borne or soil borne, or they may be spread by way of wind, water (rain, irrigation water), insects, animals, and man.
Infection by fungal pathogens depends on several factors: free water on the host plant surface is usually required, the susceptibility of the host, the density of inoculum, and ambient temperature, as well as other environmental factors. While some fungi attack only one or a few host species, others attack many. Symptoms and disease development are a function of the host-parasite interaction. Symptoms may be similar or distinct, depending on the fungi involved. Positive identification of fungi should therefore be based on their morphology. Unless otherwise reported, the fungi included in this field manual cause diseases in bread wheat, durum wheat, and triticale.
Symptoms: The postules are circular or slightly elliptical, smaller than those of stem rust, usually do not coalesce, and contain masses of orange to orange-brown urediospores. Infection sites primarily are found on the upper surfaces of leaves and leaf sheaths (1), and occasionally on the neck and awns.
Development: Primary infections usually are light and develop from wind-borne urediospores that may have travelled long distances. The disease can develop rapidly when free moisture is available and temperatures are near 20C. Successive generations of urediospores can be produced every 10-14 days if conditions are favorable. As plants mature or when environmental conditions are not favorable, masses of black teliospores may become evident (2).
Hosts/Distribution: Leaf rust can affect wheat, triticale and many other related grasses. The disease is found wherever temperate cereals are grown. The alternate hosts are Thalictrum, Isopryum, Anemonella, and Anchusa spp.
Importance: Severe early infections can cause significant yield losses, mainly by reducing the number of kernels per spike, test weights, and kernel quality.
Symptoms: Pustules (containing masses of urediospores) are dark reddish brown, and may occur on both sides of the leaves, on the stems, and on the spikes (3). With light infections the pustules are usually separate and scattered, but with heavy infections they may coalesce. Prior to pustule formation, "flecks" may appear. Before the spore masses break through the epidermis, the infection sites feel rough to the touch; as the spore masses break through, the surface tissues take on a ragged and torn appearance.
Development: Primary infections are usually light and develop from wind-borne urediospores that may have travelled long distances. The disease can develop rapidly when free moisture (rain or dew) and moderate temperatures prevail. If temperatures average about 20C or more, the first generation of urediospores will be produced in 10-15 days. As plants mature, masses of black teliospores may be produced.
Hosts/Distribution: Stem rust can affect wheat, barley, triticale, and many other related grasses; it is found wherever temperate cereals are grown. The alternate hosts are Berberis and Mahonia spp.
Importance: If infection occurs during the early crop stages, the effects can be severe: reductions in tillering and losses in grain weight and quality. Under favorable conditions, complete crop loss can occur.
Symptoms: The pustules of stripe rust, which, contain yellow to orange-yellow urediospores, usually form narrow stripes on the leaves (4). Pustules also can be found on leaf sheaths, necks, and glumes (5).
Development: Primary infections are caused by wind-borne urediospores that may have travelled long distances. The disease may develop rapidly when free moisture (rain or dew) occurs and temperatures range between 10-20C. At temperatures above 25C, the production of urediospores is reduced or ceases and black teliospores are often produced (6).
Host/Distribution: Stripe rust can attack wheat, barley, triticale, and many other related grasses. The disease is found in all highland and/or temperate areas where cereals are grown. No alternate host is known.
Importance: Severe infections can cause yield losses, mainly by reducing the number of kernels per spike, test weights, and kernel quality.
Tilletia caries, T. foetida, T. controversa
Symptoms: The main symptoms caused by these three species are fungal structures called "bunt balls," which resemble kernels but are completely filled with black teliospores. The bunt balls of common bunt, caused by T. caries and T. foetida, are about the same size and shape as the kernels they replace (7); those of dwarf bunt, caused by T. controversa, are more nearly spherical (8). When bunt balls are crushed, they give off a fetid or fishy odor. Infected spikes tend to be bluish green in color (or darker), and the glumes tend to spread apart slightly; the bunt balls often become visible after the soft dough stage (9,10; page 11). A slight reduction in plant height is typical of common bunt, while a pronounced reduction in height is typical of dwarf bunt.
Development: Spores lying dormant in the soil or on seed germinate and infect emerging seedings. Infection is favored by cool temperatures during germination. The disease develops systemically, with visible symptoms appearing after heading.
Hosts/Distribution: Wheat and (less commonly) triticale are affected by these diseases, as are several other related grasses. Common and dwarf bunts can occur worldwide. Both are limited to temperate climates; dwarf bunt occurs in areas having prolonged snow cover.
Importance: Considerable yield losses can occur when susceptible cultivars are grown or chemical seed treatments are not used.
Tilletia indica (syn. Neovossia indica)
Symptoms: Karnal bunt is not easily detected prior to harvest, since it is usual for only a few kernels per spike to be affected by the disease. Following harvest, diseased kernels can be easily detected by visual inspection: a mass of black teliospores replaces a portion of the endosperm, and the pericarp may be intact or ruptured (11). Diseased kernels give off a fetid or fishy odor when crushed.
Development: Karnal bunt is a seed- or soil-borne, floral infecting disease. inoculum (teliospores) on or near the soil surface germinates, producing sporidia, which are carried by wind to the floral structures. These sporidia in turn germinate and penetrate the glumes, rachis, or the ovary itself. The fungus enters the newly formed kernel and develops in the intercellular space between the endosperm and seed coat. The degree of disease establishment and development depends on environmental conditions from spike emergence through grain filling.
Hosts/Distribution: Karnal bunt can affect wheat, triticale, rye, and several other related grasses, but not barley. The disease is endemic in the Asian Subcontinent and now in Mexico.
Importance: Karnal bunt is a relatively minor disease. Actual losses in yield are minimal, but the disease is on the quarantine lists of many countries and therefore of importance in world grain trade.
Symptoms: The entire inflorescence, except the rachis, is replaced by masses of smut spores (12). These black teliospores often are blown away by the wind, leaving only the bare rachis and remnants of other floral structures(13).
Development: Wind blown teliospores that land on the flowers of wheat plants can germinate and infect the developing embryo of the kernel. The mycelium of the loose smut fungus remains dormant in the embryonic tissues of the kernel until the kernel begins to germinate. The mycelium then develops along with the growing point of the plant, and at flowering time replaces the floral parts of the spike with masses of black spores. Infection and disease development are favored by cool, humid conditions, which prolong the flowering period of the host plant.
Hosts/Distribution: The disease can occur wherever wheat is grown.
Importance: Yield losses depend on the number of spikes affected by the disease; incidence is usually less than one percent and rarely exceeds thirty percent of the spikes in any given location.
Symptoms: Masses of black teliospores are produced in narrow strips just beneath the epidermis of leaves, leaf sheaths and occasionally the culms. Diseased plants often are stunted, tiller profusely and the spikes may not emerge. A severe infection usually induces the leaves to roll, producing an onion-type leaf appearance. The epidermis of older diseased plants tends to shred, releasing the teliospores (14).
Development: Germinating kernels or very young seedlings are infected by germinating spores on the seed or in the soil. The disease continues to develop systemically, and the black subepidermal strips of teliospores become visible near heading. Infection is favored by low soil moisture and cool soil temperatures.
Hosts/Distribution: Bread wheats are the primary hosts of flag smut fungi, and the isolates attacking bread wheat tend to be so exclusively. There are few reports of flag smut on durum wheats and triticales. The disease is found in most winter wheat areas and in cool, fall-sown spring wheat areas.
Importance: Flag smut generally is not an economically important disease, but where present, yield losses can range from trace amounts to moderate levels (when susceptible cultivars are grown).
Erysiphe graminis f. sp. tritici
Symptoms: On all hosts, the first visible symptoms of this disease are white to pale gray, fuzzy or powdery colonies of mycelia, and conidia on the upper surfaces of leaves and leaf sheaths (especially on lower leaves) (15), and sometimes on the spikes. Older fungal tissue is yellowish gray (16). This superficial fungal material can be rubbed off easily with the fingers. Host tissue beneath the fungal material becomes chlorotic or necrotic and, with severe infections, the leaves may die. Eventually, black spherical fruiting structures (cleistothecia) may develop in the mycelia, and can be seen without magnification.
Development: The development of powdery mildew is favored by cool (15-22C), cloudy, and humid (75-100% relative humidity) conditions.
Hosts/Distribution: The fungus has a high degree of host specificity. Isolates infecting wheat do so exclusively; the same appears to be true for isolates infecting barley, oats, and rye. Further specialization exists in the form of races. Powdery mildew occurs worldwide in cool, humid, and semiarid areas where cereals are grown.
Importance: Powdery mildew can cause major yield losses if infection occurs early in the crop cycle and conditions remain favorable for development so that high infection levels are reached before heading.
Development: Initial infections tend to be on the lower leaves, progressing to the upper leaves and spikes if environmental conditions remain favorable. Cool temperatures (10-15C) and prolonged wet, cloudy weather favors the development of these diseases.
Hosts/Distribution: These are primarily diseases of wheat, but other cereals are somewhat susceptible. The diseases are limited to temperate wheat-growing areas where cool and moist conditions prevail.
Importance: Major losses can occur, through seed shrivelling and lower test weights, if these diseases reach severe levels prior to harvest.
Cochliobolus sativus (Helminthosporium sativum) Syn. Bipolaris sorokiniana, Drechslera sorokiniana
Symptoms: Lesions caused by this disease are elongated to oval in shape and are generally a dark brown color. As lesions mature, the centers often turn a light brown to tan color, surrounded by an irregular dark brown ring (21 on leaf; 22 on spike).
Development: Primary infections tend to be on the lower leaves, beginning as chlorotic flecks or spots. These infection sites enlarge, turn dark brown, and often coalesce. When the disease is severe, affected leaves or leaf sheaths may die prematurely.
Hosts/Distribution: Spot blotch affects wheat, triticale, barley, and most grasses. It is found worldwide, but is especially prevalent in more humid and higher rainfall areas.
Importance: If infection occurs early in the crop cycle and conditions remain favorable for development, complete defoliation is possible; major reductions in yield and severely shrivelled kernels will then results.
Pyrenophora trichostoma (Helminthhosporium tritici-repentis) Syn. P. tritici-repentis, Drechslera tritici-repentis
Symptoms: At first, lesions appear as tan to brown flecks, which expand into large, irregular, oval- or lens-shaped tan blotches with a yellow or chlorotic margin (23). As these spots coalesce, large blotches are formed. The development of a dark brown to black spot in the center of the lesion is characteristic of the disease. As the disease progresses, entire leaves, spikes, and even whole plants may be killed.
Development: Initial infections come from diseased crop debris in the soil, or from diseased grass hosts. Usually the lower leaves are infected first, and the disease progresses to the upper leaves and leaf sheaths if conditions are favorable. This disease develops over a wide range of temperatures and is favored by long periods (18 hours or more) of dew or rain.
Hosts/Distribution: Tan spot can affect wheat and several related grasses; triticale, barley, and rye are less frequently affected. The disease is found in the major temperate wheat-growing areas.
Importance: When severe, tan spot can cause premature death of the leaves, thereby reducing yield by lowering test weights and producing a high degree of kenel shrivelling. This disease has become more important with the increased adoption of minimum and/or no-till agronomic practices, which leave crop debris in place.
Symptoms: Small, chlorotic, oval- or elliptical-shaped lesions appear and, as they enlarge, these lesions become irregular in shape. The chlorotic borders of the lesions may become diffuse and turn light to dark brown in color (24). Lesions are difficult to distinguish from those caused by Helminthosporium spp. Infection usually starts on the lower leaves, but symptoms can be found on all plant parts.
Development: The fungus survives as conidia on seed or as mycelia within seed. sporulation on lower leaves provides inoculum that can be dispersed by wind, leading to secondary spread of the disease. Seed-borne inoculum often results in spike infections late in the crop cycle. High humidity or irrigation, as well as warmer temperatures (20 to 25C) favor infection and disease development.
Hosts/Distribution: Bread wheat and durum wheat, as well as several related grasses, are the primary hosts. The disease is common in the eastern and central areas of the Asian Subcontinent.
Importance: Alternaria leaf blight can be very severe if environmental conditions are favorable for disease development; major losses can result when susceptible cultivars are grown.
Calonectria nivalis (Fusarium nivale)
Symptoms: The blotching caused by this organism becomes evident on leaves at about late-joint to early-boot growth stage. Young lesions occur as oval to elliptical, greyish green mottled areas, usually located where the leaf bends (25). The lesions enlarge rapidly, developing into large, "eyespot" blotches with bleached or light grey centers; the leaves tend to split or shred, beginning at the centers of the lesions (26). The fungus also can cause seedling blight, foot rot, head scab (27) and, in winter cereals, pink snow mold.
Development: Spores are produced on crop debris left on or near the soil surface. These spores are transmitted to leaves by the wind or by splashing rain. Disease development is favored by cool, moist weather.
Hosts/Distribution: Generally, the disease affects durum wheat and triticale more than bread wheat or rye; oats and barley appear to be immune. Reports indicate that the disease is restricted to East Africa, the highland areas of Mexico, the Andean region of South America, and parts of southern China.
Importance: Severe disease development can cause complete defoliation, resulting in poor grain development, shrivelling, and low test weights (28).
Symptoms: Infected florets (especially the outer glumes) become slightly darkened and oily in appearance (29). Conidiospores are produced in sporodochia, which gives the spike a bright pinkish color (30). Infected kernels may be permeated with mycelia and the surface of the florets totally covered by white, matted mycelia.
Development: Several species of Fusarium can attack the spikes of small grain cereals; the ovaries are infected at anthesis, and infection is favored by warm and humid weather during and after heading. Temperatures between 10 and 28C are required for infection. Once primary infection has occurred, the disease can spread from floret to floret by mycelial growth through the spike structure.
Hosts/Distribution: All small grain cereals may be affected by this disease. Fusarium spp. are present in nearly all soils and crop residues.
Importance: Severe levels of infection can cause yield losses of more than 50% and significant reductions in grain quality. Kernels from diseased spikes are often shrivelled. Harvested grain containing more than 5% infected kernels can contain enough toxin to be harmful to humans and animals.
Symptoms: At flowering, infected florets produce a yellowish, sticky, sweet exudate (containing conidia) that is visible on the glumes. As the spike matures, kernels of infected florets are replaced by brown to purplish black fungal structures (sclerotia or "ergot bodies") (31). These ergot bodies can reach 20 mm in length(32).
Development: The primary infection originates from ascospores in fruiting bodies produced by sclerotia from the previous year's crop. Ascospores infect the florets, which then produce the sticky exudate containing conidia. Insects are attracted to the sweet exudate, and carry conidia to healthy florets in the same spike or to adjacent spikes. Rainy or humid weather favors the production of exudate and spores. An ergot body develops in each infected floret; these fungal structures can survive in the soil from one season to the next, and under dry conditions they can remain viable for many years. Sclerotia require cold temperatures before they can germinate.
Hosts/Distribution: Ergot is found in all small grain cereal crops, especially if sterility occurs for some reason (e.g., frost). Sterile florets tend to open and thus become more liable to infection. The disease is more prevalent in cool, humid climates.
Importance: Yield losses tend to be small, but losses due to discounted grain quality can be significant and occur worldwide.
Alternaria, Helminthosporium, and Fusarium spp.
Symptoms: The pericarps of maturing wheat kernels turn dark brown to black, with the discoloration usually restructed to the germ-end of the kernel (33; 34 is healthy seed). If caused by Alternaria spp., the dark color affects only the pericarp; if caused by Helminthosporium or Fusarium spp., the germ may be invaded and injured or killed. There are other fungi that can cause black point, but the three noted here are the most common.
Development: Usually, kernels are infected by these fungi during the dough stage. If humid weather prevails for a few days to a week just prior to harvest, the incidence of infection will increase and black point will develop in many cultivars.
Hosts/Distribution: Wheat is the principal host; triticale and several related grasses also can be affected. Distribution is worldwide, wherever small grain cereals are grown.
Importance: Losses are due mainly to discounted prices paid for discolored grain; if Fusarium or Helminthosporium spp. are involved, the viability of the seed also may be reduced.
Sclerophthora macrospora (Sclerospora macrospora)
Symptoms: Diseased plants tiller profusely; they have short, erect, irregular, or crooked yellowish-green culms; and the leaves are thick, erect, and usually in whorls (35). Tillers die prematurely or never head. If formed, the heads may be branched, and some of the floral tissues grow into leaflike structures (36).
Development: The disease in cereals is generally associated with water-logged or excessively irrigated fields. Development is enhanced if temperatures range between 10 and 25C. Infection may be initiated by inoculum in the soil, or from diseased weed hosts, and water must be present for infection to occur. Symptoms are most obvious during the tillering/stem elongation growth stages of the host plant.
Hosts/Distribution: Small, localized epidermics can occur when conditions are favorable. There have been no reports of widespread and destructive epidemics.
Importance: Small, localized epidemics can occur when conditions are favorable. There have been no reports of widespread and destructive epidemics.
Gaeumannomyces graminis f. sp. tritici Syn. Ophiobolus graminis
Symptoms: This fungus causes rotting of the roots and lower stems. Basal stem and leaf sheath tissues, as well as roots, may turn a shiny black color (37). When examined with a hand lens (10x), dark fungal hyphae may often be found on the subcrown internode beneath the old leaf sheaths. Coarse, black runner hyphae or conspicuous on roots. Severe disease development is indicated by stunted plants with whitened stems and spikes (38). When infection occurs early in the crop cycle, the number of tillers is often reduced and spikes are often sterile.
Development: The fungus persists on crop debris in the soil. Initial infections come from contact with hyphae or ascospores in the soil. Infection can occur throughout the crop cycle, but is favored by cool (12-18C) soil temperatures and alkaline or nutrient deficient soils. Nitrate also appears to enhance disease development. Infections of the roots occurring in the fall and early spring generally progress to the crown and lower stem tissues; infections occurring later in the crop cycle cause less damage since they usually are confined to the roots.
Hosts/Distribution: The take-all fungus displays a degree of specialization for wheat, triticale and several related grasses. The disease appears to be restricted to temperate wheat-growing areas.
Importance: Take-all is widespread in monocropped areas and has been known to cause considerable yield losses in winter wheat and fall-sown spring wheat areas, especially where liming or minimum tillage is practiced.
Pseudocercosporella herpotrichoides Syn. Cercosporella herpotrichoides
Symptoms: The most obvious symptoms of this disease are the eye-shaped, elliptical lesions produced on the internodes of the lower stem (39). The lesions are bordered by dark brown to greenish brown rings, have straw-colored centers, and frequently develop on the leaf sheath at soil level. These lesions may coalesce and lose their distinct "eye-spot" appearance. When disease development is severe, the stem or culm may break near the ground or through the lesion where the stem is weakened (40). Symptoms do not appear on the roots.
Development: Primary infection occurs from conidia or mycelia produced on crop debris on or near the soil surface; contact with the developing coleoptile or basal areas of young culms is required. The fungus is limited to the basal areas of the plant. Disease development is favored by cool, damp weather, and by high humidity at the soil level.
Hosts/Distribution: Wheat, triticale, rye, oats, and other related grasses can be affected by the disease, with wheat being the most susceptible; winter wheat and fall-sown spring wheat are more frequently damaged. Eyespot occurs in cool, moist climates where fall-sown cereals predominate.
Importance: Eyespot may kill individual tillers or even whole plants. More commonly, yield losses result from reduced kernel size and number, and from lodging.
Symptoms: The primary symptoms of sharp eyespot are the lesions that develop on basal leaf sheaths; these lesions are similar to those caused by Pseudocercosporella herpotrichoides (eyespot). Sharp eyespot lesions are more superficial and more sharply outlined than those typical of eyespot. The margins are dark brown with pale, straw-colored centers (41). The mycelia often present in the centers of lesions are easily removed by rubbing. Roots can also be affected, usually becoming brown in color and reduced in number. The disease can cause stunting and a reduction in the number of tillers.
Development: Infection is highly dependent upon environmental conditions. Disease development is favored by dry, sandy soils, cool temperatures, and high humidity. The fungus persists in soil and in crop debris, and invades root and crown tissues.
Hosts/Distribution: Rhizoctonia solani has perhaps the widest host range of any pathogen, attacking most cultivated crops; virtually all members of the Graminae family are susceptible. It is a ubiquitous fungus, present almost everywhere in soil and crop debris.
Importance: The disease is usually more severe in fields that are continously sown to cereals, especially winter wheat. However, no major or widespread epidemics have been reported.
Helminthosporium, Fusarium, and Pythium spp.
Symptoms: These fungi produce a darkening or browning of the root, crown, and basal culm tissues. Individual plants or groups of plants may lodge. White spikes often are visible just prior to normal physiological maturity (42). Infection early in the crop development can cause pre- or post-emergence "dampong off" of seedlings (43). Since each fungus can attack a different plant part at a different growth stage, positive field identification of the causal agent is difficult.
Development: The roots or crown tissues are infected by conidia or mycelia present on crop debris. Infection by and development of common root rot (Helminthosporium spp.) is favored by warm, dry soils when the plants are under stress. The root, foot, and crown rots caused by Fusarium spp. Are generally favored by cooler, moist soils, and wet soils favor Pythium spp.
Hosts/Distribution: These diseases affect all major small grain cereals grown in the temperate regions of the world.
Importance: Major epidemics have not been reported. However, localized losses can result from thinned stands, a decrease in the number of tillers, and from reductions in head size and tests weights.
Corticum rolfsii (Sclerotium rolfsii)
Symptoms: If infection occurs early in the crop cycle, pre- or post-emergence "damping off" of seedlings can result. Diseased tissues will frequently have white, fluffy fungal mycelia on the surface (44), which often permeate the soil surrounding the plant. Subsequent disease development results in rotted culms, crowns and roots, and the eventual death of the plant; this leads to the appearance of "white heads" or spikes in the green crop (45). Sclerotia are commonly found on the crown tissues, culms, or near the soil surface (46). Young sclerotia are whitish and turn brown to dark brown with age.
Development: Sclerotium rolfsii can attack the plant at any stage of development. Fungal mycelia on crop debris or sclerotia serve as primary inoculum. Infection and disease development are favored by warm (20C +) temperatures, excessive moisture, and acid soils.
Hosts/Distribution: Most cereals and grasses, plus many broad-leaf plant species are susceptible, and the fungus is widespread in tropical and subtropical environments.
Importance: The disease is seldom a problem in traditional wheat-growing areas.
Alternaria, Cladosporium, Stemphylium, Epicoccum and other species.
Symptoms: The typical symptom is the blackened appearance of mature or dead spikes, caused by a superficial accumulation of mycelia and sporulating fungal tissue (47).
Development: When wet or humid weather occurs at or near crop maturation, or when plants are heavily infested with aphids, or when plants die prematurely, they may be invaded by one or more of these fungi. Technically, black molds do not constitute a disease, since the fungi are saprophytic and invade only dead or dying plant tissue.
Hosts/Distribution: Black molds affect any kind of dead or dying plant tissue; distribution is worldwide.
Importance: Black molds generally are not economically important. Under humid or rainy conditions, the fungi can invade mature kernels, causing discoloration, black point, or smudge.
Bacterial plant pathogens are small unicellular rods from 1 to 3 æm in length. They do not have a well-defined nucleus, nor a nuclear membrane. Bacteria are spread by insects, air currents, splashing rain, and by mechanical means. Free moisture usually is necessary for infection, and penetration of host tissue occurs through wounds or stomatal openings. These pathogens invade the vascular system or intercellular spaces in host tissue, and necrosis results from toxins produced or enzymatic activity of the bacteria.
Xanthomonas campestris pv. translucens Syn. X. translucens, X. translucens flsp. undulosa, X. campestris pv. undulosa
Symptoms: Black chaff and bacterial stripe are both caused by the same organism; the site and extent of the symptoms depends on the strain of the bacterium, the affected cultivar and environmental conditions. "Bacterial black chaff" occurs primarily on the glumes (48); "bacterial stripe" occurs primarily on the leaves and/or leaf sheaths (49). Initial symptoms are narrow chlorotic lesions or stripes that have a water-soaked appearance; droplets of sticky yellowish exudate may appear with extended periods of rain or dew (50). The exudate dries to form crusty droplets or a translucent film on the surface of affected tissues. The film may crack and give a scaly appearance. If infection occurs early in the crop cycle, the spike may be infected, resulting in sterility; when the disease is severe, entire leaves or spikes may be killed.
Development: The bacterium can be seed borne and persists on crop residues in the soil, tolerating warm as well as freezing temperatures. Free moisture is required for infection and spread of the broken epidermal tissue. The disease is spread by splashing rain, plant contact, and insects.
Hosts/Distribution: These diseases occur worldwide on all small grain cereals and many grasses.
Importance: Black chaff and bacterial stripe rarely cause significant damage, even though symptoms often may be extensive.
Pseudomonas syringae pv. atrofaciens Syn. Pseudomonas atrofaciens
Symptoms: The leaves, culms, and spikes of wheat and triticale can be infected. Infections begin as small, dark green, water-soaked lesions that turn dark brown to blackish in color. On the spikelets, lesions generally start at the base of the glume and may eventually extend over the entire glume (51). Diseased glumes have a translucent appearance when held toward the light. Dark brown to black discoloration occurs with age. The disease may spread to the rachis, and lesions may also develop on the kernels (52). Under wet or humid conditions, a whitish gray bacterial ooze may be present. Stem infections result in dark discoloration of the stem; leaf infections result in small, irregular, water-soaked lesions. Symptoms can be confused with those of other bacterial diseases, genetic melanism (false black chaff), septoria nodorum blotch (glume blotch), and frost damage.
Development: The pathogens survives on crop debris, as well as various grass hosts. It is disseminated by splashing rain or by insects, and can be seed borne.
Hosts/Distribution: The disease can affect all small grain cereal crops; distribution is worldwide.
Importance: Basal glume rot usually is not economically important, but is frequently reported in humid cereal-growing areas.
Symptoms: A yellow exudate on the spikes is indicative of bacterial spike blight. When dry, the exudate is white. Often the spikes and necks will emerge as a distorted, sticky mass (53). The early leaves may also be wrinkled or twisted. This bacteria is associated with the nematode Anguina tritici in some regions.
Development: The bacteria persists in organic material in the soil. It attacks wheat when it comes in contact with the plant apex within the leaf whorl, and this transmission is often facilitated by the nematode A. tritici.
Hosts/Distribution: Wheat is the only cultivated host, though some wild grasses are susceptible to attack. The disease is frequently reported in the Asian Subcontinent.
Importance: Bacterial spike blight is not economically important.
Viruses are the smallest pathogens presented in this field guide, and only one viral disease (BYD) is discussed. The infectious viral particle is called a virion, which is a stable, non-multiplying stage by which the virus is transferred from one plant to another. Viruses multiply in the host plant, and transmission may occur via several means: by insects and mites (especially sucking insects, such as aphids), by nematodes, by seed, by pollen, by fungi, by soil and mechanically.
Viral diseases are often difficult to detect because infected hosts may not display visible symptoms, or symptoms may closely resemble those of various physiological disorders or genetic abnormalities. Identification can be facilitated by determining which vectors are present and the host range; in many cases, positive identification requires the use of an electron microscope and serological techniques.
Symptoms: The symptoms of barley yellow dwarf (BYD) vary with the affected crop cultivar, the age of the plant at the time of infection, the strain of the virus, and environmental conditions. Symptoms often are masked by or confused with other problems. Affected plants show a yellowing (54) or reddening (on oats and some wheats) of leaves, stunting, an upright posture of thickened stiff leaves (55), reduced root growth, delayed (or no) heading, and a reduction in yield. The heads of affected plants tend to remain erect and become black and discolored during ripening due to colonization by saprophytic fungi.
Development: Temperatures of approximately 20C are favorable for disease development and symptoms appear approximately 14 days after infection.
Vectors/Hosts/Distribution: The term barley yellow dwarf virus (BYDV) includes several related viruses that are all aphid transmitted. Over 20 species of aphids may act as vectors. BYD is probably the most widely distributed viral disease of cereals in the world. It attacks not only wheat, but also barley, triticale, oats, and many other grass species.
Importance: Infections occurring early in the crop cycel can result in yield losses of more than 20%, and much larger losses have been recorded.
Wheat and triticale can be attacked by a great many insects. Fortunately, only a few insect species are of major importance, causing severe damage over large geographical areas; most species are only occasional pests and/or are not geographically widespread. The "pest status" of many of species is not always well documented.
Symptoms: Aphids are nearly transparent, soft-bodied sucking insects (56). When present in sufficient numbers, aphids can cause yellowing and premature death of leaves. They exude drops of sugary liquid known as "honeydew", which may cause tiny scorch marks on the foliage and tends to encourage the development of sooty molds. The feeding of Schizaphis graminin (57) is especially damaging, resulting in the development of necrotic areas sometimes accompanied by purpling and rolling of the infested leaves. The feeding of Diuraphis noxis produces long white stripes on the leaves (58), leaf rolling, postrate growth habit, and sterile heads.
Life Cycle: The life cycle of aphids involve winged (alates), wingless (apterous), sexual, and asexual forms. When feeding on cereals, the females of most aphid species reproduce asexually (without being fertilized), giving rise to nymphs rather than eggs.
Hosts/Distribution: Species commonly found on cereals throughout the world include:
Importance: Aphids are important and widespread pests on cereal crops. When feeding in sufficient numbers, they can cause significant damage. In addition, the species listed above may act as vectors of barley yellow dwarf virus.
Symptoms: Adult stink bugs feed on stem tissue or developing kernels (59). Saliva from this insect is toxic to the plant, and a single feeding puncture can kill a stem. Feeding on kernels during the milk dough stage will destroy the kernel, while feeding during later development stages will badly shrivel the grain. Feeding on the developing head may cause partial or total sterility. Adult stink bugs have a shield-shaped body (60) and emit a disagreeable odor when crushed.
Life Cycle: Stink bugs over-winter as adults and may diapause. They tend to hibernate under dead leaves and grass. In the spring they migrate to cereal hosts, mate, and lay eggs at various places on the plant. These hatch into nymphs that feed on the plant. Mild winters and low rainfall seem to favor outbreaks of the insects.
Hosts/Distribution: Stink bugs will feed on most cereals and grasses, as well as a large range of weeds (depending on the species). Stink bugs are of major economic importance in Asia Minor.
Importance: Losses due to stink bugs are highly variable and depend on the density of the insects, weather conditions, and duration of the crop growing period. Losses are due primarily to reduced baking quality.
Symptoms: The primary symptom is defoliation of the plant. Larvae feed on leaves, chewing from the edges to the midrib, or on the heads of cereal plants. Heavy infestations can be very destructive; larvae may climb the plant and sever the neck just below the head. Some species may be found feeding at the soil surface, others underground feeding on roots, and still others feeding inside the stem.
Life Cycle: Adult cutworms (61) and army worms (62) are moths, and the females lay eggs on leaves and leaf sheaths near the ground. These eggs hatch within a few days and initially the larvae (63, cutworm; 64, armyworm) feed close to where they hatch. The larvae are found in cracks in the soil or under rocks during the day, feeding at night or early in the morning. In damp weather, they may feed all day.
Hosts/Distribution: Larvae are generally omnivorous in attacking grasses. Species of these insects are found in most cereal-growing areas of the world.
Importance: Cutworms and armyworms sporadically cause severe damage; when they do, they can devastate large areas.
Symptoms: Adult beetles are 4-5 mm long, have a black head, light brown thorax, and a shiny blue-green wing cover with parallel lines of small dots (65). Larvae are a dull to bright yellow color, but soon take on the appearance of a slimy, globular, black mass due to the mound of fecal material they produce and accumulate on their backs (66; 67). The most prominent symptom of cereal leaf beetle infestations is the distinct, longitudinal stripes on leaves (68); these stripes are produced by the feeding of adult beetles and of larvae.
Life Cycle: The insect produces one generation per year. Adults begin their feeding activity in the spring. They lay yellow eggs, either singly or in small chains, covering them with a sticky film that the soil and the adults emerge in summer. Adults overwinter underneath plant debris on the soil surface, in leaf sheaths and ears of standing maize, or under the bark of trees.
Hosts/Distribution: Cereal leaf beetles can be a problem on fall-sown cereals. Wheats with hairy leaves are affected less.
Importance: Significant yield losses can occur in winter wheat and fall-sown spring wheat. Yield losses of from 14% to more than 25% have occurred with natural infestations.
Symptoms: Thrips are small (1 mm long), brown or black insects with a tapering, segmented abdomen. They have piercing and sucking mouthparts and usually have two pairs of narrow wings. They are usually found behind the sheath of the flag leaf, feeding on the stem (69). However, leaves, stems, and heads may be attacked. Adults and nymphs both can cause damage and, if present in large numbers, may cause the tissue on which they are feeding to take on a silver coloration.
Life Cycle: Eggs are inserted into or attached to host tissue. The generation time is very short, and there may be 10 or more generations per year. Heavy rains will usually destroy a high proportion of they population.
Host/Distribution: Several thrips species live exclusively on cereals, and on forage or weed grasses.
Importance: Thrips rarely cause serious damage, and it is unusual to find infestations at such a level as to warrant control.
Symptoms: Severe infestations of Hessian flies result in stunting of the plants, thin stands, lodging, and reduced yield. Injury is caused entirely by the larvae, which suck juices from plant tissues (70). If infestation occurs during jointing, infested stems often will break prior to maturity. The Hessian fly is 3-4 mm long, has a black head and thorax, and a pinkish or yellow-brown abdomen.
Life Cycle: Adult flies emerge in the spring from pupae that have overwintered in straw or stubble. The minute, oblong eggs are reddish in color and are laid in rows on the upper sides of leaves. The eggs hatch within one week; the white, legless larvae settle behind the leaf sheaths and suck the sap of the plant. They develop into translucent, pale green, slug-like maggots. The reddish brown pupae, commonly called "flag seed" because of their resemblance to the seed of the flax plant, are oval shaped, flattened, taper to a point, and are 3-5 mm long. They are found behind leaf sheaths, usually at a node.
Hosts/Distribution: The Hessian fly is mainly a pest of wheat, but it may attack barley, rye, and other grasses. This pest has been reported in most wheat-growing areas of the world.
Importance: This is one of the most destructive insect pests on cereals. Widespread outbreaks have occurred and, in some locations (such as North Africa and the USA), the past recurs annually.
Symptoms: When young tillers are attacked in the fall or early spring, the tillers usually die; infested plants show the "white head" condition typically produced by stem-boring insects (71). The adult flies are about 6 mm in length, and pale green to yellow with dark stripes.
Life Cycle: Wheat stem maggot larvae overwinter in cereal plants or grasses (72). The females lay small white eggs, one per stem, near the sheath of the flag leaf; the larvae burrow into and consume the interior of the stem, killing the upper part of the stem and the head. There are normally three generations per year; one in the spring, one in the summer, and a third in the early autumn that overwinter as larvae.
Hosts/Distribution: In addition to wheat, host crops include rye, barley, and other grasses. There are a number of other flies in various parts of the world that attack wheat in a similar fashion and produce the same kind of damage.
Importance: In infested fields, 10-15% of plants may be injured. Damage can be severe in some years, but the insect seldom causes widespread damage. However, heavy infestations of individual wheat stands may kill a significant portion of the tillers.
Symptoms: Damage by sawflies includes premature yellowing of the head and shrivelling of the grain. The larvae girdle the stem (73) and, later in the crop cycle, lodging is common.
Life Cycle: Sawflies produce one generation per year. The larvae overwinter in the straw (74); in the spring they pupate. Adult sawflies are small, fly-like wasps and appear from late spring to midsummer. The females deposit small white eggs in the upper nodes of stems just below the heads. Upon hatching, the legless white larvae bore into the stem and tunnel downward, feeding on the pith of the stem. When they have completed their feeding, they descend further and girdle the stem base.
Hosts/Distribution: Nearly all cultivated cereals and native grasses act as hosts, although wheat is preferred. Fall-sown cereals are more commonly attacked. Wheat lines having solid or partially solid stems are much less susceptible to attack. The wheat stem sawfly is a major problem in the Mediterranean Basin.
Importance: Sawfly can cause significant damage in some years, but infestations are usually discontinuous.
Symptoms: White grubs can partially or completely sever the roots of the host plants. This causes patches of wilting and dying wheat plants (especially at the seedling stage), symptoms that could be attributed to root rots. However, when stunted patches are observed, the surrounding soil should be examined for the larvae (75). When fully grown, the largest of these larvae may be several centimeters long and nearly one centimeter thick. The larvae have three pairs of legs on their thorax (76).
Life Cycle: White grubs are the larvae of May or June beetles. Eggs are deposited in the soil and the hatched larvae feed on roots. The duration of the larval stage varies from species to species.
Hosts/Distribution: Many species of white grubs found throughout the world can attack wheat and many other plant species. Cereal crops may suffer significant damage if seeded into heavily infested grasslands.
Importance: When the roots are not completely destroyed, the plants may survive, but are stunted and fail to produce heads. However, the distribution and extent of attack is not uniform.
Symptoms: Wireworm damage is very similar to that caused by other soil-inhabiting chewing insects; the only sure means of identifying wireworms as the causal agent is to find them in association with the damaged seedlings (77). The name "wireworm" refers to the tough, wire-like appearance of the larvae. They are 20-30 mm long and are often smooth, hard, and highly polished. They have three pairs of legs (78), and their color may vary from a rich cream to shades of brown. Wireworm larvae may attack wheat as soon as the crop is seeded, eating the endosperm of the kernels and leaving only the seed coat. A common sign of woreworm attack is the wilting and/or dying of a number of adjacent plants, either in a row or patch. The stems of affected seedlings will be chewed just above the seed.
Life Cycle: Wireworms are the larvae of click beetles, of which there are many species. The adult beetles (79) lay eggs in the soil, usually in the spring, and the larvae may take several years to develop prior to pupating, depending on species. Generations overlap so that all stages and sizes of larvae may be found in the soil at the same time.
Hosts/Distribution: Many species of wireworms are found throughout the world, all of which can attack wheat. These larvae are capable of attacking many different plant species as well.
Importance: Woreworms are among the most damaging soil-infesting insects. Damage is usually most severe where wheat has been seeded after fallow or after a number of years of grass.
Symptoms: Slugs and snails (80) can feed on the endosperm of germinating seed, bite seedlings off at ground level, and graze older plants, chewing longitudinal stripes on the leaves (81). This gives the adult plant a frayed appearance. Grasshoppers (82) and crickets cause damage that is very similar to that caused by cutworms and armyworms.
Hosts/Distribution: All these insects will attack a large range of plant species, including the cereals. Distribution is worldwide.
Importance: These insects usually are localized in their distribution, but can cause a great deal of damage to individual stands of wheat.
Symptoms: Adult mites are usually less than 1 mm long, and most of the plant-inhabiting species have sucking mouth parts. Mites are not insects. Adults typically possess four pairs of legs (83), while larvae have only three pairs. However, the wheat curl mite, Eriophye tulipae (syn. Aceria tulipae), has only two pairs of legs. When present in large numbers, mites cause a silvery flecking on leaves (84). Some species may produce webs and/or may cause infested plants to be severely stunted, to head poorly, and to turn white. Individual mites are so small they they can scarcely be seen with the unaided eye. Even so, if an infested leaf is held over a piece of white paper (folded to form a trough) and tapped sharply several time, hundreds of mites will fall onto the paper and can be seen moving about.
Hosts/Distribution: Mites generally are not an important problem, with the exception of the wheat curl mite, which is a vector of wheat streak mosaic virus (WSMV).
Nematodes, also known as nemas or eelworms, are unsegmented round worms that inhabit soil and water in great numbers. Some species are parasitic on plants. All nematodes develop from eggs and pass through a succession of larval stages (usually four) prior to adulthood. Reproduction may be sexual or parthenogenic. Nematodes are dispersed in soil, running water, and plant parts. Some species have a resting stage that will withstand desiccation.
Nematode feeding reduces plant vigor and induces lesions, rots, deformation, and gall and root knots. Infested fields appear uneven, usually with distinct patches of stunted plants.
Symptoms: Distorted leaves and stems are evident prior to heading. As diseased plants approach maturity, galls are formed in the florets, replacing the kernels (85). The galls are similar in shape to the seed they replace and are dark brown in color (86). Large numbers of motile larvae are present within the galls and become active after the galls have been moistened. These nematodes can act as vectors of Corynebacterium tritici.
Development: Seed galls are dispersed along with seed during planting and harvest. In moist soil, seed galls release thousands of larvae. Wet weather favors larval movement and the infestation process. The nematode invades the crown and basal stem area, finally penetrating floral primordia. There they mature and produce large numbers of eggs. Seed galls develop in undifferentiated floral tissues.
Hosts/Distribution: The seed gall nematode parasitizes wheat, triticale, rye, and related grasses; it affects wheat primarily. It is found in the Near and Middle East, the Asian Subcontinent, Eastern Europe and occasionally, in North America.
Importance: This nematode rarely is of economic importance.
Symptoms: Cereal cyst nematodes are more readily detected on seedlings than on adult plants. The roots of infested plants develop frequent branches (87) and swellings (cysts). The cysts are off-white when young, turning dark brown as they age. Seedlings weakened by nematodes often are invaded by soil-borne pathogens, especially root and crown rots.
Development: Larvae in moist soil penetrate roots near the growing point and grow into adults. Cell enlargement, root swelling, and root branching occur as the nematodes mature. Cysts are formed as the nematodes continue to develop into egg-producing adults.
Hosts/Distribution: Most cereal crops and related species can be affected by cereal cyst nematodes. They are found in most cereal-growing regions of the world, especially in newly cultivated areas that were previously in pasture.
Importance: Significant yield losses can occur when nematode populations are high. All wheat cultivars are susceptible, but some do not support cyst formation.
Symptoms: Infestations of root knot nematodes are characterized by the formation of small knots or galls near the tips of the roots. Above ground, infested plants are stunted and chlorotic. Excessive branching of affected roots sometimes occurs (88).
Development: Root knot nematodes usually invade plants in the spring or early summer. Each root knot contains one or more females, which produce large egg masses within their saclike bodies. By midsummer the eggs are extruded and the nematodes overwinter as eggs. Usually there is one generation per year.
Hosts/Distribution: Root knot nematodes have a very wide host range, including all small grain cereals. Meloidogyne naasi appears to have specificity for cereals and grasses, and can be found wherever cereals are grown.
Importance: The damage caused by root knot nematodes depends on the number of egg masses in the soil. All cultivars of winter and spring wheat seem to be compatible hosts of the nematode. In factors as nutritional deficiencies, poor drainage, and soil-borne diseases can conceal the presence of nematodes.
Abnormal plant development may be due to physiologic or genetic disorders, nutrient deficiencies, and environmental stresses and irritants. Physiological leaf spots, blotches, and chlorosis of leaves may occur for many reasons. Some forms of chlorotic streaks, spotting, and necrosis are genetically inherited (such as chlorotic flecking and brown necrosis), resulting from chromosomal instability or certain nonviable genetic combinations (hybrid necrosis). Genotypes develop spotting and in the physical appearance of the spots.
Symptoms: When no pathogen can be identified as the cause of leaf spotting (89), the symptom may be caused by a physiological disorder or a mineral deficiency (for example, manganese deficiency).
Development: "Splotch" of winter bread wheats and durum wheats ir reported as a physiological leaf spot; the spots begin appearing during heading and increase in size and number toward the top of the plant. Spotted plants otherwise are normal. Leaf spots may also occur when cool, cloudy, and moist weather is followed by hot, sunny weather, or as a result of large fluctuations in temperature.
Hosts/Distribution: The occurrence of physiological leaf spots is related to variety and its interaction to the environment.
Importance: Usually not a serious problem; ongoing breeding efforts tend to eliminate genotypes prone to spotting.
Symptoms: Melanism occurs as brownish black to dark purple spots, streaks, or blotches on the leaf sheaths, stems (90), and/or glumes and results from a high production of melanoid pigments in some genotypes. The dark brown pigmentation usually develops on the glumes (91) and peduncles. Melanism and brown necrosis may be confused with bacterial black chaff or septoria glume blotch.
Development: The symptoms develop under certain environmental conditions, in particular with cloudy, humid weather and a high intensity of ultraviolet light (high elevations). Melanism appears to be linked with the stem rust resistance genes from "Hope" and H44.
Hosts/Distribution: Melanism occurs wherever wheat is grown, but is more pronouncee in high radiation, high humidity environments.
Importance: Generally not a serious problem; some crosses exhibit severe necrosis and yield is affected.
Symptoms: Numerous diseases cause chlorotic flecking, but "self-inflected" or genetically controlled flecking is a common occurrence in small grain cereals. The flecks may vary from small pinpoints to large blotches (92). In some cases, leaves may be a pale green color, which may be a genetic trait for low chlorophyll content.
Development: Genetic flecking or blotching may develop at any point in the crop cycle, but is more apparent at later stages of plant development (especially in spring wheats).
Hosts/Distribution: Found in many genotypes of wheat.
Importance: Genetic flecking does not necessarily make a wheat genotype unsuitable for cultivation. Several commercial wheat cultivars have high yield potential and have been released despite genetic flecking.
Some forms of spotting and necrosis result from chromosomal instability or genetic aberrations; these disorders are known as hybrid necrosis. Affected plants show differing degrees of necrosis (93), often associated with stunting or dwarf clumping. Hybrid necrosis usually is observed in early generation material (94), and affected progeny are eliminated.
Poor plant growth often can be attributed to inadequate levels of essential plant nutrients. Nitrogen, phosphorous, and potassium are used by the plant in relatively large amounts, and therefore are the nutrients that are most commonly deficient. However, micronutrient deficiencies can occur as well. Many minerals in the soil is too high. The buildup of salts in the soil, insufficient water, extreme temperatures, and poor application of pesticides also can affect the growth and yield of a crop.
Nitrogen deficient wheat appears pale green (95) and lower leaves become yellow, usually from the tip to the sheath, followed by necrosis if the deficiency persists. Nitrogen deficiency is the most common and widespread nutrient deficiency in small grains.
Phosphorus deficiency usually results in stunted plants with fewer shoots (96), if the deficiency is mild. Severe deficiency often causes pale to yellowish red leaves, starting with the lower leaves and moving from the leaf tips inward. Affected tissue may turn brown and, with severe deficiency, eventually dies. Green portions of the leaves may be bluish-green and the base of the culms purple. The development of small heads is also a common symptom.
Potassium deficiency can be difficult to detect, and yield losses can occur long before visual symptoms appear. A severe deficiency will cause the shortening of internodes, and the tips and margins of the lower leaves will become dry and scorched.
Maganese deficiency causes grayish necrotic spots or streaks to appear on the basal portion of the newest leaves. The necrotic spots may extend across the blade causing the upper portion of the leaf to kink or twist (97). Deficiency of manganese occurs most commonly in soils that matter. Oats are more sensitive than other small grain species. Foliar applications of manganese sulfate can alleviate this deficiency.
Copper deficiency symptoms include the discoloration of young leaf tips, followed by breaking and curling of the leaves (98). The plant may also produce bleached and sterile spikes. Often the spike does not emerge properly from the culm.
Symptoms: High concentration of aluminum will first reduce development of the roots, giving them a stubby appearance. They will often have a brownish color. Typical symptoms in the above ground portion of the plant are small leaves, and shortened and thickened internodes (99). It also is common for leaf tips to die and for old leaves to become yellow and brittle.
Development: This toxicity is associated with low soil pH, and it can be reduced by limiting.
Hosts/Distribution: Though many minerals can be toxic to plants, the most common toxicity affecting wheat is caused by an excess of free aluminum. Generic variability exists for aluminum tolerance within bread wheats and triticales (100).
Importance: Large areas of potentially productive land with acid soils (low pH) have toxic levels of free aluminum.
Symptoms: Salt concentrations within a field are rarely uniform; therefore, one of the first symptome indicating a salt problem is variability in crop growth within the field (barren spots are not uncommon) (101). Plants suffering from salt stress are stunted and dark blue-green in color, with tip burn and firing on the leaf margins. A soil test can rapidly confirm whether levels of salt in the soil are excessive.
Hosts/Distribution: All small grains are affected, but barley is more tolerant to high levels of salt than other small grain species.
Importance: In some areas, salt levels in the soil have limited yields for a long time; some poorly drained irrigated wheat areas are experiencing a buildup of salt that will eventually limit yields.
Symptoms: Moisture stress early in the crop cycle will stunt plants and reduce tillering and root development. Curling and rolling of leaves during midday also are symptoms of moisture stress (102). Moisture stress during the development of the spike can reduce the number of spikelets and florets, and severe stress may result in grain shrivelling. Other critical periods occur during late booting and during seed set. Severe water stress during these periods can cause complete or partial sterility.
Distribution: Moisture stress occurs to some extent each year in most rainfed environments.
Importance: Yield is often reduced without the appearance of obvious visual symptoms.
Symptoms: The effects of high temperatures often are associated with the effects of moisture stress, and the symptoms are difficult to separate (103). Moderately high temperatures increase the rate of plant development and reduce its rate of growth. The number and formation of spikelets and florets, as well as grain filling, are reduced, resulting in lower yields. The late-boot and seed-set stages are especially vulnerable and, in many areas, high temperatures are more likely to occur during these later stages of plant development. Very high temperatures will kill plants by denaturing proteins.
Importance: In many areas, the flowering to maturity period in wheat coincides with the beginning of hot, dry weather. If desiccating winds occur along with high temperatures, major reductions in yield may be experienced.
Symptoms: Phytotoxicity can result from the poor application of most pesticides. The application of such hormonal herbicides as 2,4-D too early in the crop cycle can cause leaf curling and deformed spikes (104); application near anthesis can cause sterility. Residues from the application of triazines (such as atrazine) to the crop preceeding wheat can adversely affect wheat growth (105); symptoms are bleaching of the leaves followed by necrosis (106).
Development: Damage results when chemicals are applied in excessive amounts, at the wrong growth stage, or to the wrong species.
Importance: In small grain cereals, damage is generally limited; deformations seldom cause significant losses.
Symptoms: Chlorosis of affected tissues is the common symptom. A light frost may only affect new tissue, resulting in a banding or striping on the leaves or spikes. A severe frost will kill affected tissue, which takes on a bleached white appearance (107). Sterility can result from frost occurring at flowering (108). The epidermis of the peduncle often becomes separated from underlying tissue.
Development: The freezing of plant tissue can occur at any stage of the crop cycle. Young or newly emerged tissue is the most susceptible to damage. Flowering parts are particularly sensitive.
Hosts/Distribution: All plants can suffer frost damage, and frosts can occur in most temperate wheat-growing regions.
Importance: Frost can be a serious problem if it occurs late in the crop cycle.
Alternate host A second host species required by some rusts and other organisms to complete their life cycle. Chlorosis Yellowing or whitening of normally green plant tissue. Coalesce To combine, to fuse or come together. Conidiophore A threadlike stalk upon which conidia (spores) are produced. Conidium Any asexual spore formed on a conidiophore. (pl. conidia) Diapause A period of dormancy Exudate Gel-like accumulation of spores or bacterial ooze. Foot rot Disease symptoms, such as discoloration, necrosis and decay, affecting the roots and basal portions of the plant or culm. Gall A localized proliferation of plant or parasite tissue that produces an abnormal growth or swelling, usually caused by pathogenic organisms, nematodes, or insects. Honeydew A sticky exudate (containing conidial) produced during one stage of the life cycle of Claviceps purpurea. Hypha A tubular, threadlike filament of fungal (pl. hyphae) mycelium. Immune Not affected by pathogens. Inoculum Spores or other diseased material that may cause infection. Lesion A visible area of diseased tissue on an infected plant. Mosaic A pattern of disease symptoms displaying mixed green and lighter colored patches. Mycelium A mass of hyphae that form the body of (pl. mycelia) a fungus. Necrosis Death of plant tissue, usually accompanied by discoloration. Pathogen A microorganism that causes disease. Primary Spores or fragments of a mycelium inoculum capable of initiating a disease. Pustule A spore mass developing below the epidermis, usually breaking through at maturity. Resistance Inherent capacity of a host plant to prevent or retard the development of an infectious disease. Sclerotium A dense, compact mycelial mass capable (pl. sclerotia) of remaining dormant for extended periods. Senescence The phase of plant growth that extends from full maturity to death. Spore A minute reproductive unit in fungi and lower plant forms. Sporulation The period of active spore production. Striate Displaying narrow parallel streaks or bands. Susceptible Being subject to infection or injury by a pathogen; non-immune. Symptom A visible response of a host plant to a pathogenic organism. Telium Postule containing teliospores. (pl. telia) Teliospore A thick-walled resting spore produced by rust and smut fungi. Tolerant The ability of a host plant to develop and reproduce fairly efficiently while sustaining disease. Transmission The spread of a disease agent among individual hosts. Urediospore An asexual spore of the rust fungi. Vector An organism capable of transmitting inoculum. Virulence The relative ability of a microorganism to overcome the resistance of a host. Water soaked Appearing wet, darkened, and partially transparent.