INTRODUCTION Nematodes are the most abundant group of multicellular animals on earth in terms of numbers of individuals.

Nematodes are triploblastic, unsegmented, bilaterally symmetrical, roundworms, that are transparent, colorless, usually bisexual, and worm-shaped (Vermiform), although some can become swollen (pyroform).There are at least as many nematode species as insect species. In a square meter of moderately fertile soil to 30 cm depth, there are about 50 million nematodes. Some scientists estimate that there are over 1 million kinds of nematodes, making them second only to the insects in numbers about 50 percent are small animals living in marine environments, and 25 percent live in the soil or fresh water and feed on bacteria, fungi, other decomposer organisms, small invertebrates or organic matter. Most nematodes are free-living and feed on bacteria, fungi, protozoans and other nematode (40% of the described species); many are parasites of animals (invertebrates and vertebrates (44% of the described species) and plants (15% of the described species). The largest nematode ever observed is Placentonema gigantisma, discovered in the placenta of a sperm whale. This 8 meter long nematode is said to have 32 ovaries. Most plant parasites are considerably smaller, usually measuring around 1 mm or less. Several Longidorus species exceed 10mm in length. THE HISTORY OF NEMATODES Members of the phylum Nematoda (round worms) have been in existence for an estimated one billion years, making them one of the most ancient and diverse types of animals on earth. They are thought to have evolved from simple animals some 400 million years before the "Cambrian explosion" of invertebrates able to be fossilized. The oldest known fossil nematodes are only 120-135 million years old; by then nematodes had diversified to feed on microbes, animals and plants .The oldest fossil nematodes are found in amber and are commonly associated with insects. This is probably due to the fact that tree sap, which fossilizes to make amber, captures and preserves insects and their associated nematodes much more easily than an animal- or a nematode-infested portion of a plant. The two nematode classes, the Chromadorea and Enoplea, have diverged so long ago, over 550 million years, that it is difficult to accurately know the age of the two lineages of the phylum (Figure 1). Phylum

Nematoda
Chromadorea Rhabditida Figure 1. Enoplea Dorylaimida Triplonchida

Classes Orders

Phytoparasitic nematodes were known in antiquity (235 B.C.) because the ancient Chinese symbol for a soybean root-infesting organism resembles in shape an adult female soybean cyst nematode

(Noel, 1992). The first described plant parasitic nematodes were discovered in wheat seeds by Needham (1743). Not until the identification of root-knot nematodes on cucumber by Berkeley (1855) and cyst nematodes causing beet-tired disease on sugar beets by Schacht (1859), did plant nematology begin to emerge as an important scientific discipline. Nathan A. Cobb, the father of US nematology, pioneered agricultural nematology as a USDA scientist in the early 1900s. The use of soil fumigation to reduce nematode populations and increase crop yields in the 1940s (Carter) demonstrated that nematodes were significant crop pathogens and ushered in the chemical era for nematode management in production agriculture. MORPHOLOGY AND ANATOMY Plant nematodes are tiny worms usually 0.25 mm to 3 mm long (1/100 "to 1 / 8 ") and cylindrical, tapering toward the head and tail. Females of a few species lose their worm shape as they mature, becoming pear-, lemon- or kidney- shaped. Plant parasitic nematodes possess all of the major organ systems of higher animals except respiratory and circulatory systems. The body is covered by a transparent cuticle, which bears surface marks helpful for identifying nematode species. LIFE CYCLE AND REPRODUCTION The life cycle of a plant-parasitic nematode has six stages: egg, four juvenile stages and adult. Male and female nematodes occur in most species, but reproduction without males is common, and some species are hermaphroditic (A females@ produce both sperm and eggs). Egg production by the individual completes the cycle. Most species produce between 50 and 500 eggs per female, depending on the nematode species and their environment, but some can produce more than 1,000 eggs. The length of the life cycle varies considerably, depending on nematode species, host plant, and the temperature of the habitat. During summer months when soil temperatures are 80 to 90 F, many plant nematodes complete their life cycle in about four weeks.

NEMATODE FEEDING AND HOST-PARASITE RELATIONSHIPS Plant parasitic nematodes feed on living plant tissues, using an oral stylet, a spearing device somewhat like a hypodermic needle, to puncture host cells. Many, probably all, plant nematodes inject enzymes into a host cell before feeding to partially digest the cell contents before they are sucked into the gut. Most of the injury that nematodes cause plants is related in some way to the feeding process. Ectoparasitic Nematodes feed on plant tissues from outside the plant; endoparasitic nematodes feed inside the tissues. If the adult female moves freely through the 61 soil or plant tissues, the species is said to be as migratory. Species in which the adult females become swollen and permanently immobile in one place in or on a root are termed as sedentary. Migratory endoparasitic and ectoparasitic nematodes generally deposit their eggs singly as they are produced, wherever the female happens to be in the soil or plant. Sedentary nematodes such as root-knot (Meliodogyne spp.), cyst (Heterodera spp.), reniform (Rotylenchulus spp.), and citrus (Tylenchulus semipenetrans) nematodes produce large numbers of eggs, which remain in their bodies or accumulate in masses attached to their bodies. The feeding/living relationships that nematodes have with their hosts affect sampling methods and the success of management practices. Ectoparasitic nematodes, which never enter roots, may be recovered only from soil samples. Endoparasitic nematodes often are detected most easily in samples of the tissues in which they feed and live (burrowing and lesion nematodes), but some occur more commonly as migratory stages in the soil (root-knot and reinform nematodes). Endoparasitic Nematodes inside root tissues may be protected from those kinds of pesticides that do not penetrate into roots. Root tissues may also shield them from many microorganisms that attack nematodes in the soil. Ectoparasites are more exposed to pesticides and natural control agents in

the soil. Foliar nematodes (Aphelenchoides spp.) are migratory nematodes that feed on or inside the leaves and buds of ferns, strawberries, chrysanthemums and many other ornamentals. They cause distortion or death of buds, leaf distortion, or yellow to dark-brown lesions between major veins of leaves. Other nematodes that attack plants above ground, cause leaf or seed galls. Still others cause deterioration of the bulbs and necks of onions and their relatives. Summary of Plant Parasitic Nematode Feeding Strategies Feeding Strategy Ectoparasite Example Genera Order Belonolaimus Xiphenema Trichodorus SemiEndoparasites Migratory Endoparasites Sedentary Endoparasites Stem Seed Nematodes Foliar Nematodes Aphelenchoides Rhabditida J2-adult Adult * eggs, all juvenile stages and adults can survive the winter, but not egg producing females SYMPTOMS CAUSED BY NEMATODES Determining if nematodes are involved in a plant growth problem is difficult because few nematodes cause distinctive diagnostic symptoms. A sound diagnosis should be based on as many as possible of: symptoms above and below ground, field history, and laboratory assay of soil and/or plant samples. Above-Ground Symptoms. It is rare that above-ground symptoms give sufficient evidence to diagnose a nematode problem in the roots. However, they are important because they are almost always the reason that nematode problems are first noticed. Since most plant nematodes affect root functions, most symptoms associated with them are the result of inadequate water supply or mineral nutrition to the tops: chlorosis (yellowing) or other abnormal coloration of foliage, stunted top growth, failure to respond normally to fertilizers, small or sparse foliage, a tendency to wilt more readily than healthy plants, and slower recovery from wilting. and Rotylenchulus Tylenchulus Pratylenchus Radopholus Meloidogyne Heterodera Naccobus Nematodes Ditylenchus Gall Anguina Rhabditida Dorylaimida Rhabditida Rhabditida Rhabditida Rhabditida Rhabditida Rhabditida Rhabditida Rhabditida Rhabditida J2 J2 J2 J4 J4 J2 J3 J4 J2 J4 vectored by insects Egg/cyst Infective Stage J2-adult J2-adult J4 J2 J2-adult J4 J2 * Vector viruses Vector viruses Resistant Notes Stage

Triplonchida J2-adult

Bulb Bursaphelenchus Rhabditida

Woody plants in advanced stages of decline caused by nematodes may exhibit dieback of progressively larger branches. A Melting out, or gradual decline, is typical of nematode- injured turf and pasture. Plantings stunted by nematodes often have worse weed problems than areas without them because the crop is less able than it should be to compete with weeds. Distribution. The distribution of nematodes within any site is very irregular, so the shape, size and distribution of areas with the most severe effects of nematodes will be erratic within the field. Nematodes move very few feet per year on their own. In the undisturbed soil of groves, turf and pastures, visible symptoms of nematode injury normally appear as round, oval or irregular areas that gradually increase in size year by year. In cultivated land, nematode injured spots are often elongated in the direction of cultivation because nematodes are moved by machinery. Erosion, land leveling, and any other force that moves masses of soil or plant parts can also spread a nematode infestation much more rapidly than it will go by itself. Nematode damage is often seen first and most pronounced in areas under special stresses , such as heavy traffic, excessive drainage because of slope or soil and dry areas outside regular irrigation patterns. Below-Ground Symptoms may be more useful than top symptoms for diagnosing nematode problems. Galls caused on roots by root-knot nematodes, abbreviated 62 roots or stunted root growth, necrotic lesions in the root cortex, and root rotting may all be symptoms of nematode problems. An experienced observer can often see cyst nematodes (Heterodera, Globodera and Cactodera spp.) on the roots of their hosts without magnification. The young adult females are visible as tiny white beads, about the size of a pin head. After a female cyst nematode dies, her white body wall is tanned to a tough brown capsule containing several hundred eggs. Important cyst nematodes found in Florida include soybean cyst nematode (H. glycines) on soybeans and a few leguminous weeds, beet cyst nematode (H. schachtii) on cabbage and related plants, St. Augustine grass cyst nematode (H. leuceilyma) on St. Augustine grass and cactus cyst nematode (C. cacti) on Christmas cactus and related plants. H. cyperi is a cyst nematode occasionally found infesting nutsedges (Cyperus spp). DIAGNOSING NEMATODE Field History. Accurate field history can provide valuable clues to the identity of nematode and other pest problems. A nematode that has been present in the field in recent years is probably there yet, and is likely to injure susceptible crops if environmental conditions are favorable. Production records that show a gradual decline in yields over a period of years despite no change in cultural practices may indicate progressive development of a nematode problem. A nematode infestation in a new field usually begins in a small area. It gradually intensifies in the original spot and is spread through the field by cultivation, harvest, erosion and other factors that spread infested soil or plant parts. Therefore, the total effect of a recently introduced nematode is a gradual production decline for the field, as the percentage of the field that is involved and the severity of damage at any given area in the field increase over the

years. Laboratory Assay. Laboratory analysis of soil and/or plant tissue samples is often necessary to complete a diagnosis. In the lab, nematodes are extracted from soil and plant tissues, identified, and counted. Those results can be compared with research and field observations to determine whether or not the crop is likely to be injured by the population under those conditions. In some cases, specific steps to reduce the numbers and/or effects of a particular nematode species are recommended only if the population density exceeds some predetermined level felt to represent the threshold for economic loss of that crop. Such thresholds are determined through longterm experience of nematologists with that pest and crop in growers operations and in controlled experiments. NEMATODE DISSEMINATION. While nematodes are motile animals, most are able to move no more than a meter through the soil within their lifetime. However, this lack of long distance crawling does not mean nematodes cannot rapidly spread from field to field. Farm equipment and even muddy shoes contaminated with nematode-infested soil can rapidly disperse nematodes. The movement of water during floods and irrigation can disperse nematodes over long distances. Likewise the movement of nematode infected plants, seeds, and bulbs can give nematodes international tickets to travel the world if plant quarantine officials are not careful. The ability of nematodes to form environmentally resistant stages makes their dissemination even easier, since dried nematodes can be blown with the wind or plant debris over large geographical regions. Even migrating birds are suspected to be able to carry nematodes along their flight paths, assisting the nematodes in their quest for new homes. Essentially any process that moves soil or plant tissue has the ability to disperse plant nematodes, making them difficult plant pathogens to quarantine. NEMATODE PESTS OF MUSHROOM AND THEIR CONTROL In practice, mushroom growers can come across such mushroom culture pests as nematodes. Two species, Ditylenchus myceliophaguss and Aphelenchoides composicola were identified by researchers as mushroom pests that disturb mycelium growth. They are characterized by a needlelike mouthpart with which they pierce the mycelial hyphae, inject digestive juices and suck out the cell contents. The cells die, and the nematodes that move from cell to cell destroy the hyphal structures. Moreover, it is assumed that nematodes that pierce mycelial threads, spread bacterial and virus diseases. In areas where mycelium dies, the compost becomes soggy and depressed an objectionable odor occurs because of anaerobic bacteria activities. These microscopic, colorless worms that live under different conditions in nature: in soil, water and decaying remains of plants. Nematodes can enter the farm in different ways: with compost, casing soil, water, insects and personnel. The nematodes found in mushroom culture are divided into two groups: Saprophytic, which feed on decaying materials Mycophytic nematodes or Parasitic, which only feed on fungi.

SAPROPHYTIC NEMATODES Saprophytic nematodes are not considered to be mushroom pests. With their oral cavity that looks like a tube, they suck in organic nutrients and microorganisms and especially bacteria that exist in great amount in compost and in the casing layer. That is why poorly prepared and pasteurized compost is not well suited for the development of mushroom mycelium, but is a favorable environment for the nutrition and reproduction of nematodes that have survived pasteurization. In nonselective substrate, with overly humid areas, nematode population grows incredibly fast. The toxins they release and the increasing number of putrefactive bacteria weaken the mycelium growth. The same thing can happen when water leaks through the casing layer after watering, which causes the mycelium to suffocate. The compost becomes watery and darkens while theres an objectionable odor in the room because of putrefactive bacteria. When the infection is severe, the nematodes swarm in great amounts on the casing surface. They form columns that look like erect waving sticks and thin threads that flicker in the light. In this condition, nematodes are able to adhere to flies, mites and personnel that harvest mushrooms.

The swarming of nematodes of the casing surface Moreover, nematodes can have the ability to dry slowly and form a cyst; a temporary protective layer due to which they can survive in adverse conditions, being in a dormant state for a long period of time. In this state, nematodes are easily spread by wind. Parthenogenesis, which is the kind of sexual reproduction that allows females to reproduce without males, makes it possible for the nematode population to grow in favorable conditions.

Half-empty beds the result of severe infection of the casing layer by nematodes

A poor harvesting technique: that is when mushroom stipes and remains are left on the beds promotes the development of nematodes, hence they will feed on decaying mushroom tissue. MYCOPHYTIC NEMATODES These nematodes feed only on fungi. Two species, Ditylenchus myceliophaguss and Aphelenchoides composicola were identified by researchers as mushroom pests that disturb mycelium growth. They are characterized by a needlelike mouthpart with which they pierce the mycelial hyphae, inject digestive juices and suck out the cell contents. The cells die, and the nematodes that move from cell to cell destroy the hyphal structures. Moreover, it is assumed that nematodes that pierce mycelial threads, spread bacterial and virus diseases. In areas where mycelium dies, the compost becomes soggy and depressed an objectionable odor occurs because of anaerobic bacteria activities. Mycophytic nematodes differ from saprophytic nematodes because they cant reproduce asexually and dont form flickering columns. The presence of nematodes on mushroom beds indicates bad cultivation conditions, that is, poor substrate and casing soil preparation and nonobservance of the rules of sanitation and hygiene. The degree of crop losses depends on the time and level of the initial infection. Infection at spawning time can have an effect on mycelial growth and thus making cultivation unprofitable from an economical point of view. Infection occurring later causes only minor crop losses and goes unnoticed by the mushroom grower. MUSHROOM NEMATODES CONTROL MEASURES The nematode control measures are mainly used to prevent their spreading on the farm. This can be achieved, if all operations are carried out properly. Well prepared and pasteurized compost ensures fast and healthy mycelial growth and makes it impossible for the pests to spread. When its suspected that the source of infection is peat (though the bottom layers of peat in beds dont contain pests), all incoming batches of peat should be tested for the presence of nematodes. Peat that contains pathogenic organisms has to be disinfected and stored in a clean room in order to escape recurring infection.

Growing rooms that are infected with nematodes (if its rational) should be kept at low temperatures and dry, as far as it is possible. In the end of the cultivation cycle, they have to be steamed, making sure the temperature remains at the same level in all parts of the room 70C. When steaming is impossible, the compost is fumigated with methyl bromide. All spent compost must be immediately removed from the farm territory. Measures must be taken to prevent the spread of nematodes by insects, personnel and implements. At normal hygienic conditions and short cultivation cycles (2-3 breaks), nematodes dont often cause serious problems. However, mushroom growers must use all resources in order to prevent the occurrence and spreading of nematodes on the farm, hence its impossible to control them during the cultivation cycle.

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Identification of nematodes: Nematodes feeding: SCN mating: Nematodes movies: Phylogenetic trees of nematodes: http://nematode.unl.edu/key/nemakey.htm http://www.cropsci.uiuc.edu/faculty/lambert-k/ http://www.cropsci.uiuc.edu/faculty/lambert-k/ http://www.cropsci.uiuc.edu/faculty/lambert-k/ http://www.wormbook.org/chapters/www_quicktour diversity/ quicktourdiversity.html

University of Nebraska Lincoln http://nematode.unl.edu/ Nematology: Iowa State University, Soybean cyst http://nematode.unl.edu/scn/scnisu.htm nematode: Alternative nematode control: Soybean cyst nematode: Nematode identification: The virtual nematode: Other nematodes: UC Davis Nematode databases: Society of nematologists: C. elegans information: Biocontrol web site: History of Nematology: http://www.attra.org/attra-pub/nematode.html http://extension.missouri.edu/xplor/agguides/crops/g 04450.htm http://kbn.ifas.ufl.edu/gaster/identify.htm http://www.ppws.vt.edu/~sforza/nematode.html http://elegans.swmed.edu/Nematodes/ http://ucdnema.ucdavis.edu/search.html http://www.nematologists.org/ http://elegans.swmed.edu/ http://sacs.cpes.peachnet.edu/nemabc/ http://flnem.ifas.ufl.edu/nemadoc.htm

Chemical methods of nematode control: http://plpnemweb.ucdavis.edu/nemaplex/Mangmnt/C hemical.htm Mushroom growing http://agaricus.ru/en/doc/show/378/

http://www.apsnet.org/edcenter/intropp/PathogenGroups/Pages/IntroNematodes.aspx http://www.ext.colostate.edu/pubs/crops/02952.html http://www.fao.org/DOCREP/006/Y4011E/y4011e0p.htm