Aphids Aphids, also known as plant lice (and in Britain as greenflies), are small planteating insects, and members

of the superfamily Aphidoidea. Aphids are among the most destructive insect pests on cultivated plants in temperate regions. The damage they do to plants has made them enemies of farmers and gardeners the world over, but from a purely zoological standpoint they are a very successful group of organisms. About 4,400 species of 10 families are known. Historically, many fewer families were recognized, as most species were included in the family Aphididae. Around 250 species are serious pests for agriculture and forestry as well as an annoyance for gardeners. They vary in length from one to ten millimetres. Natural enemies include predatory lady

beetles (Coleoptera: Coccinellidae), hoverfly wasps, aphid midge larvae, crab

larvae (Diptera: Syrphidae), parasitic (Neuroptera: Chrysopidae),

spiders lacewings

and entomopathogenic fungi like Lecanicillium lecanii and theEntomophthorales. Aphids are distributed worldwide, but are most common in temperate zones. Also, in contrast to many taxa, species diversity is much lower in the tropics than in the temperate zones. They can migrate great distances, mainly through passive dispersal by riding on winds. For example, the currant lettuce aphid (Nasonovia ribisnigri Mosley) is believed to have spread from New Zealand to Tasmania in this way.Aphids have also been spread by human transportation of infested plant materials. Taxonomy Aphids are in the superfamily Aphidoidea in the homopterous division of the order Hemiptera. Recent classification within Hemiptera has reduced the old taxon "Homoptera" to two suborders: Sternorrhyncha (e.g., aphids, whiteflies, scales, psyllids, etc.) and Auchenorrhyncha(e.g., cicadas, leafhoppers, treehoppers, planthoppers, etc.) with the suborder Heteroptera containing a large group of insects known as thetrue bugs. More recent reclassifications have substantially rearranged the families within Aphidoidea: some old families were reduced to subfamily rank (e.g., Eriosomatidae), and many old subfamilies elevated to family rank. Taxonomically woolly conifer

aphids like the pine aphid, the spruce aphid and the balsam woolly aphid are not true aphids, but adelgids, and lack the cornicles of true aphids. Relation to phylloxera and adelgids Aphids, adelgids, and phylloxerids are very closely related, and are either placed in the insect super family Aphidoidea (Blackman and Eastop, 1994) or into two super families (Phylloxeroidea and Aphidoidea) within the order Homoptera, the plant-sucking bugs. Like aphids, phylloxera feed on the roots, leaves and shoots of grape plants, but unlike aphids do not produce honeydew or cornicle secretions. Phylloxera

(Daktulosphaira vitifoliae), are insects which caused the Great French Wine Blight that devastated European viticulture in the 19th century. Similarly, adelgids also feed on plant phloem. Adelgids are sometimes described as aphids, but more properly as classified as aphid-like insects, because they have no cauda or cornicles. Anatomy Most aphids have soft, green bodies, but other colors are common, such as black, brown, and pink. Aphids have antennas with as many as six segments. Aphids feed themselves through sucking mouthparts called stylets, enclosed in a sheath called a rostrum, which is formed from modifications of the mandible and maxilla of the insect mouthparts. They have long, thin legs and two-jointed, two-clawed tarsi. Most aphids have a pair of cornicles (or "siphunculi"), abdominal tubes through which they exude droplets of a quick-hardening defensive fluid containing

triacylglycerols, called cornicle wax. Other defensive compounds can also be produced by some types of aphids.

Aphids have a tail-like protrustion called a "cauda" above their rectal apertures. They have two compound eyes, and an ocular tubercle behind and above each eye, made up of three lenses (called triommatidia). When host plant quality becomes poor or conditions become crowded, some aphid species produce winged offspring, "alates", that can disperse to other food sources. The mouthparts or eyes are smaller or missing in some species and forms. Diet Many aphid species are monophagous (that is, they feed on only one plant species). Others, like the green peach aphid (Myzus persicae), feed on hundreds of plant species across many families. Aphids passively feed on sap of phloem vessels in plants, as do many of their fellow members of Hemiptera such as scale insects and cicadas. Once a phloem vessel is punctured, the sap, which is under high pressure, is forced into the aphid's food canal. As they feed, aphids often transmit plant viruses to the plants, such as to potatoes, cereals, sugarbeets and citrus plants. These viruses can sometimes kill the plants. Plants contain low densities of the nitrogen compounds needed for building proteins. This requires aphids to consume an excess of sap to satisfy their nutritional requirements. The excess is expelled as "honeydew", out of the recta of aphids, in such large volumes that in some instances it can "fall like rain". Aphid honeydew is rich in carbohydrates, like the phloem it derives from. Mutualism Some species of ants "farm" aphids, protecting them on the plants they eat, eating the honeydew that the aphids release from the terminations of their alimentary canals. This is a "mutualistic relationship".

These "dairying ants" "milk" the aphids by stroking them with their antennae. Therefore, sometimes aphids are called "ant cows". Some farming ant species gather and store the aphid eggs in their nests over the winter. In the spring, the ants carry the newly-hatched aphids back to the plants. Some species of dairying ants (such as the European yellow meadow ant, Lasius flavus) manage large "herds" of aphids that feed on roots of plants in the ant colony. Queens that are leaving to start a new colony take an aphid egg to found a new herd of underground aphids in the new colony. These farming ants protect the aphids by fighting off aphid predators. An interesting variation in ant-aphid relationships involves Lycaenid butterflies (such as the Sievers blue butterfly and the Japanese copper butterfly) and the Myrmica ants. For example, Niphanda fusca butterflies lay eggs on plants where ants tend herds of aphids. The eggs hatch as caterpillars which feed on the aphids. The ants do not defend the aphids from the caterpillars, but carry the caterpillars to their nest. In the nest, the ants feed the caterpillars, which produce honeydew for the ants. When the caterpillars reach full size, they crawl to the colony entrance and form cocoons. After two weeks, butterflies emerge and take flight. Some bees in coniferous forests also collect aphid honeydew to make "forest honey". Many aphids are host to endosymbiont bacteria, Buchnera, inside specialized cells called bacteriocytes. These bacteria synthesize some essential amino acids that are absent from the phloem that the aphids eat. Evolution Aphids probably appeared around 280 million years ago, in the early Permian period. They probably fed on plants like Cordaitales or Cycadophyta. The oldest known aphid fossil is of the species Triassoaphis cubitus Evans from the Triassic. The number of species was small, but increased considerably with the appearance of angiosperms

160 million years ago. Angiosperms allowed aphids to specialize. Organs like the cornicles did not appear until the Cretaceous period. Threats Aphids are soft-bodied, and have a wide variety of insect predators. Aphids also are often infected by bacteria, viruses and fungi. Aphids are affected by the weather, such as precipitation, temperature and wind. Insects that attack aphids include predatory lady bugs (Coleoptera:

Coccinellidae) (ladybirds in the UK), hoverfly larvae (Diptera: Syrphidae), parasitic wasps, aphid midge larvae, aphid lions, crab spiders and lacewings (Neuroptera: Chrysopidae). Fungi that attack aphids include Neozygites fresenii, Entomophthora, Beauveria bassiana, Metarhizium anisopliae and entomopathogenic fungi like Lecanicillium lecanii. Aphids brush against the microscopic spores. These spores stick to the aphid, germinate and penetrate the aphid's skin. The fungus grows in the aphid hemolymph (i.e., the counterpart of blood for aphids). After about 3 days, the aphid dies and the fungus releases more spores into the air. Infected aphids are covered with a woolly mass that progressively grows thicker until the aphid is obscured. Often the visible fungus is not the type of fungus that killed the aphid, but a secondary fungus. Aphids can be easily killed by unfavorable weather, such as late spring freezes. Excessive heat kills the symbiotic bacteria that some aphids depend on, which makes the aphids infertile. Rain prevents winged aphids from dispersing, and knocks aphids off plants and thus kills them from the impact or by starvation. However, Ken Ostlie, an entomologist with the University of Minnesota Extension Service, suggests that rain should not be relied on for aphid control.

Defenses Aphids are soft-bodied, and have little protection from predators and diseases. Some species of aphid interact with plant tissues forming a gall, an abnormal swelling of plant tissue. Aphids can live inside the gall, which provides protection from predators and the elements. A number of galling aphid species are known to produce specialised "soldier" forms, sterile nymphs with defensive features which defend the gall from invasion. For example, Alexander's horned aphids are a type of soldier aphid that has a hard exoskeleton and pincer-like mouthparts. Infestation of a variety of Chinese trees by Chinese sumac aphids (Melaphis chinensis Bell) can create a "Chinese gall" which is valued as a commercial product. As "Galla Chinensis", Chinese galls are used in Chinese medicine to treat coughs, diarrhea, night sweats, dysentry and to stop intestinal and uterine bleeding. Chinese galls are also an important source of tannins. Some species of aphid, known as "woolly aphids" (Eriosomatinae), excrete a "fluffy wax coating" for protection. The cabbage aphid (Brevicoryne brassicae) stores and releases chemicals that produce a violent chemical reaction and strong mustard oil smell to repel predators. It was common at one time to suggest that the cornicles were the source of the honeydew, and this was even included in the Shorter Oxford English Dictionary and the 2008 edition of the World Book Encyclopedia.. In fact, honeydew secretions are produced from the anus of the aphid, while cornicles mostly produce defensive chemicals such as waxes. There also is evidence of cornicle wax attracting aphid predators in some cases. Aphids are also known to defend themselves from attack by parasitoid wasps by kicking. Effects on plants Plants exhibiting aphid damage can have a variety of symptoms, such as decreased growth rates, mottled leaves, yellowing, stunted growth, curled leaves, browning, wilting, low yields and death. The removal of sap creates a lack of vigour in

the plant, and aphid saliva is toxic to plants. Aphids frequently transmit disease-causing organisms like plant viruses to their hosts. The green peach aphid (Myzus persicae) is a vector for more than 110 plant viruses. Cotton aphids (Aphis gossypii) often infect sugarcane, papaya and groundnuts with viruses. Aphids contributed to the spread of late blight (Phytophthora infestans) among potatoes in the Great Irish Potato Famine of the 1840s. The cherry aphid or black cherry aphid, Myzus cerasi, is responsible for some leaf curl of cherry trees. This can easily be distinguished from 'leaf curl' caused by Taphrina fungus species due to the presence of aphids beneath the leaves. The coating of plants with honeydew can contribute to the spread of fungi which can damage plants. Honeydew produced by aphids has been observed to reduce the effectiveness of fungicides as well. A hypothesis that insect feeding may improve plant fitness was floated in the mid-1970s by Owen and Wiegert. It was felt that the excess honeydew would nourish soil micro-organisms, including nitrogen fixers. In a nitrogen poor environment, this could provide an advantage to an infested plant over a noninfested plant. However, this does not appear to be supported by the observational evidence. The damage of plants, and in particular commercial crops, has resulted in large amounts of resources and efforts being spent attempting to control the activities of aphids. Banana Bunchy Top Banana bunchy top disease is caused by a virus called banana bunchy top virus (BBTV). The disease was recorded as causing serious damage to bananas in Fiji Islands as early as 1889. The virus is widespread across the Pacific Islands region, but distribution is patchy. It is present on some islands, absent on others. By late 2004, laboratory test records confirming presence of BBTV in Fiji Islands, Tonga, Samoa and New Caledonia had been published. There are also unpublished laboratory test records

of BBTV in Guam and on Wallis Island (but not Futuna). In addition there are reliable reports, dating back many years, of distinctive symptoms of the disease seen in the field in Tuvalu and American Samoa. These records are not known to have beeen confirmed in a diagnostic test. SYMPTOMS Plants in an advanced stage of infection are stunted and the throat of the plant is choked with a rosette of short, narrow, erect leaves, giving the typical bunchy top appearance. These leaves are brittle and snap off crisply when broken. They also have yellow margins that may eventually turn brown and appear burnt. The root system of such plants is poor and decayed. When a plant becomes infected, the first and second leaf produced after infection may also have a yellowish margin. The third leaf to emerge after infection is reduced in size, more yellowish and has wavy leaf edges that curve upward At this stage, closer examination can reveal further symptoms that help with preliminary field diagnosis of the disease. However, in the case of critical diagnoses such as new outbreaks in new areas, these must be confirmed properly in a laboratory. If the second or third leaf is held up to the light and examined from the underside a dot dash Morse code like series of dark green lines can be seen running parallel to the veins. These dark green lines continue into the midrib as distinct hooks . Vein clearing may also occur. Dark green streaks can often be found on the back of the petioles. Papaya ringspot potyvirus It is a devastating virus disease. In Africa, it occurs in Kenya, Nigeria, Tanzania and Uganda. Initially, the disease appears as oil streaks on stems and petioles and as it progresses, mottling of leaves becomes evident. Severely infected plants do not flower and die young. Infected fruits develop characteristic line patterns, which form rings and

remain green when fruits ripen. The virus is spread by aphids and it is also mechanically transmitted. Peanut stunt virus Peanut stunt virus may cause disease in peanut, tobacco, clover, soybean and snapbean. This relatively new virus was first found in Virginia and North Carolina in 1964, when it caused severe loss in crop yield and value. Following early infection, a plant may never grow beyond a few inches in height and width. Leaves from plants infected with peanut stunt virus are malformed and curl up at the edges. Infected leaves may be paler green and/or yellowed.

The fruit of plants infected with the virus is frequently small, malformed and the shells are commonly split open to expose seed. Infected peanut seed do not play a role in the spread of the disease, since only seed too small for planting are infected at a high enough rate to act as a source of infection. The virus can overwinter in wild or forage legumes (clovers, alfalfa, lespedeza, etc.) and then spread to other crops in the spring by aphids that carry the virus in their mouthparts after feeding on infected plants. Peanut yield and value are reduced by peanut stunt virus because of a decrease in numbers of fancy pods, extra large kernels, and sound mature Sugarcane Mosaic Virus Disease The disease caused by sugarcane mosaic virus (SCMV) is commonly referred to as "mosaic." It has, at one time or another, occurred in virtually every important sugarcanegrowing country. Estimated yield losses due to the disease vary greatly depending on the time period and sugarcane-growing area involved. Historically, it has been a serious disease problem in Louisiana. In fact, mosaic, superimposed on already established diseases in Louisiana, caused a near collapse of the industry in the mid-1920s. Until 1996, mosaic had not been a problem in Florida. In 1996, sugarcane mosaic was observed in grower fields on CP 72-2086, a major commercial cultivar. The epicenter of the disease was near where Hatton Highway intersects with US 98. Presently some fields of CP 72-2086 are infested with SCMV; the source of infection appears to be

infected seedcane. Only traces of mosaic have been found in other cultivars. The western region of the south Florida sugarcane growing area still remains essentially free of mosaic. SYMPTOMS Mosaic is identified primarily by its leaf symptoms. As with most sugarcane diseases, the symptoms may vary in intensity with the cane variety, growing conditions, and the strain of the virus involved. The most distinctive symptom is a pattern of contrasting shades of green, often islands of normal green on a background of paler green or yellowish chlorotic areas on the leaf blade. Generally, the chlorotic areas are diffuse, but they may be sharply defined in some clones infected with certain strains of the virus. The infection may be accompanied by varying degrees of leaf reddening or necrosis. Chlorotic areas are most evident at the base of the leaf. Chlorotic areas may also be present on the leaf sheath, but rarely on the stalk. Young, rapidly growing plants are more susceptible to infection than more mature, slower growing plants. Cowpea mosaic virus CPMV) is a plant virus of the comovirus group. Infection of a susceptible cowpea leaf results in high virus yields (1-2 g/kg). Its Genome consists of 2 molecules of positivesense RNA (RNA-1 and RNA-2) which are separately encapsidated. Both RNA1 and RNA2 have a VPg (virus genome-linked protein) at the 5'end, and polyadenylation at the 3' end. Genomic RNA1 and RNA2 are expressed by a polyprotein processing strategy. RNA1 encodes helicase, VPg, protease and RdRp. RNA2 encodes movement protein and coat protein. The virus particles are 28 nm in diameter and contain 60 copies each of a Large (L) and Small (S) coat protein. The structure is well characterised to atomic resolution, and the viral particles are thermostable. CPMV displays a number of features that can be exploited for nanoscale biomaterial fabrication. Its genetic, biological and physical properties are well characterised, and it can be isolated readily from plants. There are many stable mutants already prepared that allow specific modification of the capsid surface. It is possible to attach a number of

different chemicals to the virus surface[1][2] and to construct multilayer arrays of such nanoparticles on solid surfaces. This gives the natural or genetically engineered nanoparticles a range of properties which could be useful in nanotechnological applications such as biosensors, catalysis and nanoelectronic devices. One example use of CPMV particles is to amplify signals in microarray based sensors. In this application, the virus particles separate the fluorescent dyes used for signaling in order to prevent the formation of non-fluorescent dimers that act as quenchers.[3] Another example is the use of CPMV as a nanoscale breadboard for molecular electronics. Tobacco Mosaic Virus 1. The first plant virus discovered, tobacco mosaic virus (TMV), attacks members of the nightshade, or solanaceae, family. These include tobacco, pepper, potato, tomato, eggplant, cucumber and petunia. The virus spreads through entry into breaks of cell walls caused by insects or other physical damage. It has another claim to fame as the first virus imaged with an electron microscope. Cucumber Mosaic Virus 2. Cucumber mosaic virus infects cucumber, tomato, peppers, melons, squash, spinach, celery, beet and other plants. Aphids spread it, and they cause physical damage to the plant, which allows entry of the virus via wind, splashing or dripping sap. The virus causes twisting in young leaves. That stunts growth of the entire plant and causes poor fruit or leaf production. Barley Yellow Dwarf 3. The barley yellow dwarf virus infects several grains and staple crops, including wheat. Aphids primarily spread the virus. The virus causes discoloration of leaves and the tips of the plants, which reduces photosynthesis, stunts growth and decreases production of seed grains.

Bud Blight 4. The bud blight virus infects soybeans, a staple crop. It causes the stem to bend at the top and the buds to turn brown and drop off the plant. Nematodes spread this virus. Sugarcane Mosaic Virus 5. The sugarcane mosaic virus discolors leaves of the sugarcane plant, restricting its ability to feed itself through photosynthesis and grow. It stunts the growth of young plants. Aphids and infected seed spread the virus. Cauliflower Mosaic Virus 6. The cauliflower mosaic virus infects members of the brassica, or mustard, family, which includes cabbage, brussels sprouts, cauliflower, broccoli and rape seed. It causes a mosaic or mottle on the leaves, which stunts growth. Aphids and mechanical exposure spread the virus. Lettuce Mosaic Virus 7. The lettuce mosaic virus mottles the leaves of almost all types of lettuce, stunting its growth and eliminating its market appeal. Aphids and infected seeds spread the virus. Maise Mosaic Virus 8. The maise mosaic virus causes yellow spots and stripes on the leaves of corn, stunting its growth. Leafhoppers spread the virus. Peanut Stunt Virus
9. The peanut stunt virus causes discoloration and distortion of the leaves of

peanuts and some other rhizomes, stunting their growth. Aphids and sap spread the virus.

Reference: Bar-Joseph, M.; Marcus, R.; Lee, R.F. (1989) The continuous challenge of citrus tristeza virus control. Annual Review of Phytopathology 27, 291-316. Bar-Joseph, M.; Roistacher, C.N.; Garnsey, S.M. (1983) The epidemiology and control of citrus tristeza disease. In: Plant virus epidemiology (Ed. by Plumb, R.T.; Thresh, J.M.), pp. 61-72. Blackwell Scientific Publications, Oxford, UK. Bov, J.M.; Vogel, R. (Editors) (1981) Description and illustration of virus and viruslike diseases of citrus. Setco-IRFA, Paris, France. Candresse, T.; Martelli, G.P. (1995) Closterovirus genus. Archives of Virology, Supplement 10, 461-464. CMI (1978) Distribution Maps of Plant Diseases No. 289 (edition 5). CAB International, Wallingford, UK. Costa, A.S.; Mller, G.W. (1980) Tristeza control by cross-protection. Plant Disease 64, 538-541. Dodds, J.A.; Jordan, R.L.; Roistacher,C.N.; Jarupat, T. (1987) Diversity of citrus tristeza virus isolates indicated by dsRNA analysis. Intervirology 27, 177-188.