33:
1600:; Altincicek, Boran; Anselme, Caroline; Atamian, Hagop; Barribeau, Seth M; de Vos, Martin; Duncan, Elizabeth J; Evans, Jay D; Gabaldón, Toni; Ghanim, Murad; Heddi, Adelaziz; Kaloshian, Isgouhi; Latorre, Amparo; Moya, Andres; Nakabachi, Atsushi; Parker, Benjamin J; Pérez-Brocal, Vincente; Pignatelli, Miguel; Rahbé, Yvan; Ramsey, John S; Spragg, Chelsea J; Tamames, Javier; Tamarit, Daniel; Tamborindeguy, Cecilia; Vincent-Monegat, Caroline; Vilcinskas, Andreas (2010).
51:
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however pressure to maintain these pathway genes was reduced due to redundancy as a result of the presence of the other partner's genome. Unlike other related insects, the A. pisum genome is missing genes necessary for the urea cycle. the purine salvage pathway, and other genes that code enzymes necessary for the biosynthesis of molecules. These missing reaction intermediates are likely provided by genes within the
952:(broad bean). The phloem saps of these plants are nutritionally rich in carbohydrates but poor in terms of nitrogen. The ratio of essential amino acids to nonessential amino acids in these phloem saps ranges from 1:4-1:20. This ratio of essential to nonessential amino acids is severely disproportional compared to the 1:1 ratio present in animal tissues and necessary for survival. Animals, including
516:
756:- As most phloem feeders, the pea aphid is adapted to feeding on a limited set of plants. Studies on pea aphids have identified candidate loci, molecular and physiological mechanisms that are involved in host nutrition and virulence. Genetic, molecular and physiological studies have also evidenced specialization to different host species as a motor of
778:
1013:
sheath cells. There are about 60-80 bacteriocytes in each pea aphid and are organized into the bi-lobed bacteriome. A bacteriome is a specialized organ that runs along the length of the pea aphid on two sides of the body and joins near the hindgut. Bacteriocytes are located near the ovariole cluster and
1012:
cell has an inner and outer gram-negative cell membrane and is individually enclosed in an aphid-derived symbiosomal membrane. These encased cells are then grouped into specialized, aphid-derived bacteriocytes (mycetocytes). Bacteriocytes are large, polyploid cells surrounded by a thin lining of flat
356:
exist in pea aphids. Besides differences between sexual and parthenogenetic morphs, winged and wingless morphs exist. Overcrowding and poor food quality may trigger the development of winged individuals in subsequent generations. Winged aphids can then colonize other host plants. Pea aphids also show
1073:
genome has undergone more unique genomic changes compared to other insects of the order
Hemiptera. The aphid genome is 464MB with aphid-specific orphan genes making up 20% of the genome and gene duplication present in more than 2000 gene families. These orphan genes and gene duplications are likely
1179:
genome lacks IMS, dFADD, Dredd and Retish genes that are a part of the IMD (immunodeficiency) pathway and present in other related insects. Also missing are peptidoglycan recognition proteins (PGRPs) that detect pathogens and alert the IMD pathway as well as antimicrobial peptide (AMP) genes which
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are incomplete. Jointly, the genomes of these two organisms complement each other to produce complete metabolic pathways for the biosynthesis of nutrients such as amino acids and other essential molecules. The ancestral partners of this symbiosis are likely to have had complete metabolic pathways,
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contribute to the metabolic pathways of amino acid biosynthesis. This metabolic complementarity is illustrated by the use of asparagine, a nonessential amino acid in phloem sap, as a major precursor in the production essential and nonessential amino acids necessary for the growth and survival of
2194:
Brinza, Lilia; Viñuelas, José; Cottret, Ludovic; Calevro, Federica; Rahbé, Yvan; Febvay, Gérard; Duport, Gabrielle; Colella, Stefano; Rabatel, Andréane; Gautier, Christian; Fayard, Jean-Michel; Sagot, Marie-France; Charles, Hubert (2009). "Systemic analysis of the symbiotic function of
Buchnera
1065:
genome is missing genes required for surface membrane construction such as lipopolysaccharides and phospholipids as well as genes associated with cellular defense. Transporter genes and regulatory genes are also missing from the genome. Such gene loss is typical of an obligate and intracellular
530:
is considered as the model aphid species. Its reproductive cycle, including the sexual phase and the overwintering of eggs, can be easily completed on host plants under laboratory conditions, and the relatively large size of individuals facilitates physiological studies. In 2010, the
725:) made by pea aphids, is one of few carotenoids known to be synthesized by animals. Torulene imparts natural, red-colored patches to some aphids, which possibly aid in their camouflage and escape from predation by wasps. The aphids have gained the ability to synthesize torulene by
416:
is considered among the aphid species of major agronomical importance. Yields can be affected by the sap intake that directly weakens plants, although pea aphids seldom reach densities that might significantly reduce crop production. However, like many aphid species,
2576:
Wilson, A. C. C.; Ashton, P. D.; Calevro, F.; Charles, H.; Colella, S.; Febvay, G.; Jander, G.; Kushlan, P. F.; MacDonald, S. J.; Schwartz, J. F.; Thomas, G. H.; Douglas, A. E. (2010). "Genomic insight into the amino acid relations of the pea aphid,
498:
during the 1870s, and by 1900 it had become a serious pest species in the mid-Atlantic states. By the 1950s, it was widespread throughout the United States and Canada. Its host range in North
America is very similar to that of the closely related
1045:
genomes have experienced unique modifications that are likely related to the establishment and maintenance of the endosymbiotic relationship. The genomes of both organisms have undergone significant gene loss compared to related organisms. The
960:
but cannot synthesize nine essential amino acids that must be obtained through their diets: histidine, isoleucine, leucine, lysine, methionine, phenylalanine, threonine, tryptophan, and valine. In addition to these nine essential amino acids,
1240:
1078:
specific gene duplications of amino acid transporters highly expressed in bacteriocytes have been observed. These duplications are likely associated with the genetic establishment and maintenance of the endosymbiotic relationship.
1036:
were the first insect-endosymbiont pair to have the genomes of both partners sequenced. This has provided researchers with a great deal of information about the evolutionary and molecular interactions of this endosymbiosis. The
306:, like most aphids. Each adult female gives birth to four to 12 female nymphs per day, around a hundred in her lifetime. These develop into mature females in about seven to ten days. The life span of an adult is about 30 days.
688:- Pea aphid lineages include parthenogenesis in their life cycles, and some have even lost the sexual phase. Pea aphids are models for deciphering the origin and consequences of asexual reproduction, an important question in
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1319:
It is particularly important on peas, beans, alfalfa and clover, but also attacks beets, cucurbits, various species of
Brassicaceae. It has been implicated in the transmission of over 40 plant viruses.
709:
and physiological mechanisms underlying body color, reproductive cycle and the presence of wings in males (which is genetically based) have been identified in pea aphids or are being investigated.
397:. Unlike many aphid species, pea aphids do not tend to form dense colonies where individuals would stay where they were born during their whole lifetimes. Pea aphids are not known to be farmed by
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are produced once the immune pathway has been activated. A reduced immune system may have facilitated the establishment and sustained maintenance of the symbiotic relationship between the
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have been shown to actively take up 14C labeled glutamine (a nonessential amino acid) where it is then converted into glutamic acid. This glutamic acid is then taken up by the individual
1530:
Thomas, Cyrus (1877). "A List of the
Species of the Tribe Aphidini, family Aphidae, found in the United States, which have been heretofore named, with descriptions of some New Species".
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is the only species with a sequenced genome known to be missing key components of the purine salvage pathway, essential for the production of DNA, RNA, signaling molecules, and ATP. The
796:
2904:
Ramsey, J. S.; MacDonald, S. J.; Jander, G.; Nakabachi, A.; Thomas, G. H.; Douglas, A. E. (2010). "Genomic evidence for complementary purine metabolism in the pea aphid,
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Sandström, J.; Pettersson, J. (1994). "Amino acid composition of phloem sap and the relation to intraspecific variation in pea aphid (Acyrthosiphon pisum) performance".
3139:
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cells and used to synthesize the essential amino acids isoleucine, leucine, phenylalanine, and valine as well as nonessential amino acids that can be returned to
1669:
Tsuchida, T.; Koga, R.; Horikawa, M.; Tsunoda, T.; Maoka, T.; Matsumoto, S.; Simon, J. -C.; Fukatsu, T. (2010). "Symbiotic
Bacterium Modifies Aphid Body Color".
3178:
2734:
Sasaki, Tetsuhiko; Ishikawa, Hajime (1995). "Production of essential amino acids from glutamate by mycetocyte symbionts of the pea aphid, Acyrthosiphon pisum".
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genome has retained genes required for the biosynthesis of essential amino acids but has not retained genes responsible for the degradation of amino acids. The
2772:
Wilson, Alex C. C.; Sunnucks, Paul; Hales, Dinah F. (2003). "Heritable genetic variation and potential for adaptive evolution in asexual aphids (Aphidoidea)".
981:. This nutritional provisioning has been examined genomically (metabolic complementary, discussed below) and experimentally. Isolated bacteriocytes containing
349:
is avoided by the recognition of close kin. Mating between close kin has significantly lower egg hatching success and offspring survival than outbred mating.
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genome is 641-kb and consists of a circular chromosome with 2 plasmids. It has been reduced to one-seventh of the size of its closest free-living relative,
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When the colony begins to become overcrowded, some winged females are produced. These disperse to infest other plants, where they continue to reproduce
2005:
Mutti, N. S.; Louis, J.; Pappan, L. K.; Pappan, K.; Begum, K.; Chen, M. -S.; Park, Y.; Dittmer, N.; Marshall, J.; Reese, J. C.; Reeck, G. R. (2008).
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848:. This is an obligate, symbiotic relationship and both partners are completely dependent on each other. When treated with antibiotics to remove the
2360:
Baumann, P.; et al. (1995). "Genetics, Physiology, and
Evolutionary Relationships of the Genus Buchnera - Intracellular Symbionts of Aphids".
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genome. Through this complementation, the nucleotide requirements of both organisms are fulfilled: the purine salvage pathway is completed for
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insect to have been published. The pea aphid genome and other of its features are the focus of studies covering the following areas:
814:
339:. When temperatures become colder and day lengths shorter, sexual winged females and males appear. These mate, the females lay
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probably resulted from the introduction of some of its host plants for agriculture. Such an introduction likely occurred into
1810:
532:
3157:
1954:
Hawthorne, D. J.; Via, S. (2001). "Genetic linkage of ecological specialization and reproductive isolation in pea aphids".
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eggs and the life cycle starts again. Pea aphids can complete their whole reproductive cycle without shifting host plant.
317:. In autumn, the lengthening of the night triggers the production of a single generation of sexual individuals (males and
3269:
50:
1903:
Nancy A. Moran; Tyler Jarvik (2010). "Lateral
Transfer of Genes from Fungi Underlies Carotenoid Production in Aphids".
3022:
3009:
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are known to host pea aphids, though the complete host range remains undetermined. On crops such as peas and alfalfa,
2318:
Douglas, A. E. (1998). "Nutritional
Interactions in Insect-Microbial Symbioses: Aphids and Their Symbiotic Bacteria
3092:
2858:"Genome Expansion and Differential Expression of Amino Acid Transporters at the Aphid/Buchnera Symbiotic Interface"
2007:"A protein from the salivary glands of the pea aphid, Acyrthosiphon pisum, is essential in feeding on a host plant"
235:(and colloquially known as the green dolphin, pea louse, and clover louse), is a sap-sucking insect in the family
3183:
3105:
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endosymbiotic relationship is likely to have evolved 160-280 million years ago. Phylogenetic analysis shows that
1188:. Also, phloem sap is a diet with reduced amounts of microbes which may have lower the evolutionary pressure of
3014:
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has lost genes that would allow it to live outside the host but maintains genes essential for the nutrition of
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to overcome this lack of essential amino acids in the phloem sap When provided with nonessential amino acids,
726:
619:
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genome on the other hand, contains 66 amino acid biosynthesis genes and 93 amino acid degradation genes. Both
475:, which are specialized on different host species. Therefore, the pea aphid is more accurately described as a
2987:
1219:
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associated with the “metabolic, structural and developmental” components of the endosymbiotic relationship.
592:
is transmitted from mothers to offspring, and it has coevolved with aphids for dozens of millions of years.
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It was previously believed that lateral gene transfer was responsible for the severe gene reduction in the
2949:
1763:
Tsuchida, T.; Koga, R.; Fukatsu, T. (2004). "Host Plant
Specialization Governed by Facultative Symbiont".
900:
infection of the common ancestor of aphids and co-speciation of the holosymbiont has occurred since then.
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holosymbiont is one of the most well studied symbiotic relationships both genetically and experimentally.
741:- The pea aphid genome presents high levels of gene duplication compared to other insect genomes, such as
696:
442:
32:
860:
lacks genes required for living independent of a host and is unculturable outside of the aphid host. The
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1285:(134 ed.). Urbana, Illinois, USA: University of Illinois Agricultural Experiment Station. pp.
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942:
757:
660:
453:
422:
155:
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Douglas, A. E. (1993). "The nutritional quality of phloem sap utilized by natural aphid populations".
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Scarborough, C. L.; Ferrari, J.; Godfray, H. (2005). "Aphid Protected from Pathogen by Endosymbiont".
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is unable to synthesize arginine due to missing urea cycle genes. The endosymbiotic relationship with
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In the autumn, female pea aphids lay fertilized eggs overwinter that hatch the following spring. The
1418:(Hemiptera : Aphididae). Bulletin of the British Museum (Natural History) Entomology 26 (1971).
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cells are transferred to eggs during oogenesis or to the developing embryos during embryogenesis.
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cells are vertically transferred from the mother's ovaries through transovarial transmission. The
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523:- this red morph shows the reddish/dark markings due to carotenoids that some individuals produce.
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Sandström, Jonas; Moran, Nancy (1999). "How nutritionally imbalanced is phloem sap for aphids?".
1987:
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Dixon, A. F. G. Aphid Ecology - An optimization approach. 2 edn, (Kluwer Academic Pub, 1998).
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Mutual nutrient provisioning is likely the main reason for the persistence of this symbiosis.
623:
622:, and which affect ecologically important traits in aphids, such as body color, resistance to
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272:
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published an annotated draft sequence of the pea aphid genome composed of approximately 525
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diseases to the plants it visits. Protection against pea aphids includes the use of chemical
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utilizes an endosymbiotic bacterium to overcome the nutritional deficiencies of phloem sap.
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genome contains the necessary genes to encode the reaction intermediates missing from the
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evolved from a bacterium that originally occupied the gut of the aphid common ancestor.
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components controlling the production of sexual and winged morphs, among other features.
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2649:"Genomic revelations of a mutualism: the pea aphid and its obligate bacterial symbiont"
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2006:
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shows that the genome lacks expected genes essential to immune response pathways. The
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329:
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132:
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2288:"A molecular clock in endosymbiotic bacteria is calibrated using the insect hosts".
1940:
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2241:"Genome sequence of the endocellular bacterial symbiont of aphids Buchnera sp. APS"
1991:
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Population densities are at their highest in early summer, then decrease through
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Douglas, A. E. (2006). "Phloem-sap feeding by animals: Problems and solutions".
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converts nonessential amino acids into essential amino acids to be returned to
385: in) in adults. It generally feeds on the lower sides of leaves, buds and
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are housed in specialized, aphid-derived cells located in the hemocoel of the
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1811:"Evolutionary and functional insights into reproductive strategies of aphids"
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aphidicola, the primary endosymbiont of the pea aphid Acyrthosiphon pisum".
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676:) - Studies on pea aphids have helped to establish the environmental and
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2966:
2475:"Symbiotic bacteria enable insect to use a nutritionally inadequate diet"
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Proceedings of the Royal Society of London. Series B: Biological Sciences
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participates in an obligate endosymbiotic relationship with the bacteria
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677:
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van Emden, H. & Harrington, R. Aphids As Crop Pests. (CABI, 2007).
520:
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332:, from which new parthenogenetic females will emerge in early spring.
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1975:
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of a number of genes for carotenoid synthesis, apparently from fungi.
409:
268:
248:
92:
72:
2943:
1433:"Inbreeding Avoidance by Recognition of Close Kin in the Pea Aphid,
717:. Plants, fungi, and microorganisms can synthesize carotenoids, but
3079:
40:
Adult parthenogenetic pea aphid and progeny feeding on a pea plant
543:
514:
426:
260:
1489:
Braendle, C.; Davis, G. K.; Brisson, J. A.; Stern, D. L. (2006).
244:
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that hatch from these eggs are all females, which undergo four
771:
398:
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genome but sequencing has shown that this has not occurred.
375:
is a rather large aphid whose body can reach 4 millimetres (
263:
species of major agronomical importance. The pea aphid is a
2066:"The pea aphid complex as a model of ecological speciation"
1850:"Rarity of Males in Pea Aphids Results in Mutational Decay"
452:
Pea aphids, although collectively designated by the single
721:(3',4'-didehydro-β,γ-carotene, specifically a hydrocarbon
713:
is notable as one of few animals identified to synthesize
1241:"Noxious and beneficial insects of the state of Illinois"
2479:
Proceedings of the Royal Society B: Biological Sciences
792:
672:(the production of several discrete morphs by the same
596:
also hosts a range of facultative bacterial symbionts (
239:. It feeds on several species of legumes (plant family
2132:"Genome Sequence of the Pea Aphid Acyrthosiphon pisum"
1350:"Genome Sequence of the Pea Aphid Acyrthosiphon pisum"
1329:
1327:
361:
body color variations of green or red/pink. The green
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origin, but it is now commonly found worldwide under
321:
females) by the same parthenogenetic parent females.
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The International Aphid Genomics Consortium (2010).
888:
is a monophyletic group and that the phylogenies of
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growth and reproduction are interrupted or reduced.
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is the primary endosymbiont. Together they form the
2956:
1082:No lateral gene transfer has been detected between
896:coincide. Therefore, there was likely one original
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may be too technical for most readers to understand
747:, with the notable expansion of some gene families.
1563:. Gulf Professional Publishing. pp. 317–320.
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2130:International Aphid Genomics Consortium (2010).
2011:Proceedings of the National Academy of Sciences
1192:to maintain the immune response pathway genes.
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1809:Simon, J. C.; Stoeckel, S.; Tagu, D. (2010).
1414:Eastop, V. F. Keys for the identification of
1268:(276 ed.). US Department of Agriculture.
1218:. As such these are also promising potential
928:feeds on phloem sap of host plants including
243:) worldwide, including forage crops, such as
8:
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2123:
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1602:"Immunity and other defenses in pea aphids,
1431:Huang, Ming H.; Caillaud, Marina C. (2012).
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1265:The pea aphis with relation to forage crops
1245:Report of the State Entomologist (Illinois)
920:Like other insects of the order Hemiptera,
904:is related to Enterobacteriaceae including
872:Evolution of the endosymbiotic relationship
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31:
20:
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2774:Biological Journal of the Linnean Society
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815:Learn how and when to remove this message
799:, without removing the technical details.
588:and is necessary for aphid reproduction.
550:. This constitutes the first genome of a
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1802:
1532:Illinois Natural History Survey Bulletin
1410:
1408:
1406:
1404:
1102:Individually, the metabolic pathways of
2524:Entomologia Experimentalis et Applicata
2473:Akman GĂĽndĂĽz, E.; Douglas, A.E (2009).
1848:Brisson, J. A.; Nuzhdin, S. V. (2008).
1231:
956:, can produce nonessential amino acids
533:International Aphid Genomics Consortium
365:are generally more frequent in natural
299:. They will then begin to reproduce by
2799:Gerardo, N.M.; Wilson, A.C.C. (2011).
2647:Shigenobu, S.; Wilson, A.C.C. (2011).
1343:
1341:
1339:
1282:The insect pests of clover and alfalfa
797:make it understandable to non-experts
7:
3194:d7e1fbf8-f846-4f18-952c-579b1a804cec
3106:9dc1195f-43d9-4666-954d-73593f3b3f79
3015:Acyrthosiphon_(Acyrthosiphon)_pisum
2856:Price, D.R.G.; et al. (2011).
2239:Shigenobu, S.; et al. (2000).
519:Adult, parthenogentic pea aphid on
2417:10.1111/j.1365-2311.1993.tb01076.x
2064:Peccoud, J.; Simon, J.-C. (2010).
482:The pea aphid is thought to be of
449:, as opposed to many aphid pests.
14:
3260:Insect vectors of plant pathogens
3255:Insect pests of ornamental plants
3250:Insect pests of temperate forests
751:Interaction with host plants and
437:, and the selection of resistant
2926:10.1111/j.1365-2583.2009.00945.x
2826:10.1111/j.1365-294x.2011.05103.x
2786:10.1046/j.1095-8312.2003.00176.x
2599:10.1111/j.1365-2583.2009.00942.x
2544:10.1046/j.1570-7458.1999.00485.x
2091:10.1111/j.1365-2311.2009.01147.x
1198:Pests, diseases, and biocontrols
776:
765:Endosymbiotic relationship with
633:, and nutrition. (Specifically,
49:
2581:, with its symbiotic bacterium
2908:, and its symbiotic bacterium
2437:Journal of Experimental Botany
1135:receives necessary guanosine.
1:
2801:"The power of paired genomes"
2374:10.1146/annurev.micro.49.1.55
2756:10.1016/0022-1910(94)00080-Z
2736:Journal of Insect Physiology
2718:10.1016/0022-1910(94)90133-3
2698:Journal of Insect Physiology
2336:10.1146/annurev.ento.43.1.17
2149:10.1371/journal.pbio.1000313
1367:10.1371/journal.pbio.1000313
2324:Annual Review of Entomology
1560:Handbook of Vegetable Pests
1491:"Wing dimorphism in aphids"
760:between pea aphid biotypes.
655:decreases mortality due to
584:, which provides essential
283:Generalities and life cycle
267:for biological study whose
3291:
3275:Taxa named by Moses Harris
2209:10.1016/j.crvi.2009.09.007
1827:10.1016/j.crvi.2010.03.003
1557:Capinera, John L. (2001).
1279:Folsom, Justus W. (1909).
699:and physiology explaining
645:retard the development of
614:) that can be transmitted
3265:Insects described in 1776
3245:Agricultural pest insects
2665:10.1007/s00018-011-0645-2
1441:Journal of Insect Science
1098:Metabolic complementarity
184:
179:
161:
154:
46:Scientific classification
44:
39:
30:
23:
2914:Insect Molecular Biology
2587:Insect Molecular Biology
2197:Comptes Rendus Biologies
1815:Comptes Rendus Biologies
1629:10.1186/gb-2010-11-2-r21
1544:10.21900/j.inhs.v1i2.448
1291:pea aphid green dolphin.
1209:and the fungal pathogen
727:horizontal gene transfer
433:, natural predators and
325:sexual females will lay
231:, commonly known as the
2296:(1337): 167–171. 1993.
2032:10.1073/pnas.0708958105
1925:10.1126/science.1187113
1874:10.1126/science.1147919
1777:10.1126/science.1094611
1734:10.1126/science.1120180
1691:10.1126/science.1195463
2491:10.1098/rspb.2008.1476
2302:10.1098/rspb.1993.0098
1508:10.1038/sj.hdy.6800863
1239:Cyrus, Thomas (1879).
997:Holosymbiont structure
908:and it is likely that
524:
443:insecticide resistance
389:of legumes, ingesting
259:, and ranks among the
2875:10.1093/molbev/msr140
2397:Ecological Entomology
2070:Ecological Entomology
1304:"Acyrthosiphon pisum"
1171:Genome sequencing of
1115:genome. For example,
916:Nutritional symbiosis
758:ecological speciation
518:
3101:Fauna Europaea (new)
1454:10.1673/031.012.3901
1262:Davis, John (1915).
703:variations in aphids
690:evolutionary biology
685:Asexual reproduction
652:Regiella insecticola
636:Hamiltonella defensa
611:Regiella insecticola
599:Hamiltonella defensa
505:Acyrthosiphon kondoi
459:, encompass several
408:More than 20 legume
3270:Hemiptera of Africa
3217:acyrthosiphon-pisum
3002:Acyrthosiphon_pisum
2988:Acyrthosiphon pisum
2958:Acyrthosiphon pisum
2910:Buchnera aphidicola
2906:Acyrthosiphon pisum
2817:2011MolEc..20.2038G
2748:1995JInsP..41...41S
2710:1994JInsP..40..947S
2583:Buchnera aphidicola
2579:Acyrthosiphon pisum
2536:1999EEApp..91..203S
2409:1993EcoEn..18...31D
2257:2000Natur.407...81S
2082:2010EcoEn..35S.119J
2023:2008PNAS..105.9965M
1968:2001Natur.412..904H
1917:2010Sci...328..624M
1866:2008Sci...319...58B
1683:2010Sci...330.1102T
1604:Acyrthosiphon pisum
1435:Acyrthosiphon pisum
1205:faces threats from
833:Buchnera aphidicola
767:Buchnera aphidicola
642:Serratia symbiotica
605:Serratia symbiotica
581:Buchnera aphidicola
373:Acyrthosiphon pisum
228:Acyrthosiphon pisum
214:A. pisum destructor
165:Acyrthosiphon pisum
25:Acyrthosiphon pisum
16:Species of true bug
2449:10.1093/jxb/erj067
2362:Annu Rev Microbiol
1008:body cavity. Each
946:(red clover), and
943:Trifolium pretense
575:hosts the primary
525:
501:blue alfalfa aphid
3227:
3226:
3202:Open Tree of Life
2950:Taxon identifiers
2868:(11): 3113–3126.
2811:(10): 2038–2040.
2653:Cell Mol Life Sci
2485:(1658): 987–991.
2203:(11): 1034–1049.
1911:(5978): 624–627.
1598:Gerardo, Nicole M
1570:978-0-12-158861-8
1025:Genome sequencing
906:Escheriachia coli
825:
824:
817:
736:and expansion of
488:temperate climate
445:is documented in
224:
223:
218:
209:
200:
192:
3282:
3220:
3219:
3210:
3209:
3197:
3196:
3187:
3186:
3174:
3173:
3161:
3160:
3148:
3147:
3135:
3134:
3122:
3121:
3109:
3108:
3096:
3095:
3083:
3082:
3070:
3069:
3057:
3056:
3044:
3043:
3031:
3030:
3018:
3017:
3005:
3004:
2992:
2991:
2990:
2977:
2976:
2975:
2945:
2938:
2937:
2901:
2888:
2887:
2877:
2853:
2847:
2846:
2828:
2796:
2790:
2789:
2769:
2760:
2759:
2731:
2722:
2721:
2693:
2687:
2686:
2676:
2659:(8): 1297–1309.
2644:
2611:
2610:
2573:
2548:
2547:
2519:
2513:
2512:
2502:
2470:
2461:
2460:
2432:
2421:
2420:
2392:
2386:
2385:
2357:
2348:
2347:
2315:
2306:
2305:
2285:
2279:
2278:
2268:
2266:10.1038/35024074
2236:
2221:
2220:
2191:
2172:
2171:
2161:
2151:
2127:
2096:
2095:
2093:
2061:
2055:
2054:
2044:
2034:
2002:
1996:
1995:
1976:10.1038/35091062
1951:
1945:
1944:
1900:
1894:
1893:
1845:
1839:
1838:
1806:
1797:
1796:
1760:
1754:
1753:
1717:
1711:
1710:
1677:(6007): 1102–4.
1666:
1660:
1659:
1641:
1631:
1594:
1575:
1574:
1554:
1548:
1547:
1527:
1521:
1520:
1510:
1486:
1477:
1476:
1466:
1456:
1428:
1419:
1412:
1399:
1396:
1390:
1389:
1379:
1369:
1345:
1334:
1331:
1322:
1321:
1316:
1314:
1300:
1294:
1293:
1276:
1270:
1269:
1259:
1253:
1252:
1236:
1207:parasitoid wasps
840:is the host and
820:
813:
809:
806:
800:
780:
779:
772:
734:Gene duplication
647:parasitoid wasps
490:. The spread of
384:
383:
379:
295:before reaching
216:
207:
205:A. pisum spartii
198:
190:
167:
54:
53:
35:
21:
3290:
3289:
3285:
3284:
3283:
3281:
3280:
3279:
3230:
3229:
3228:
3223:
3215:
3213:
3205:
3200:
3192:
3190:
3182:
3177:
3169:
3164:
3156:
3151:
3143:
3138:
3130:
3125:
3117:
3112:
3104:
3099:
3091:
3086:
3078:
3073:
3065:
3060:
3052:
3047:
3039:
3034:
3026:
3021:
3013:
3008:
3000:
2995:
2986:
2985:
2980:
2971:
2970:
2965:
2952:
2942:
2941:
2903:
2902:
2891:
2855:
2854:
2850:
2798:
2797:
2793:
2771:
2770:
2763:
2733:
2732:
2725:
2704:(11): 947–955.
2695:
2694:
2690:
2646:
2645:
2614:
2575:
2574:
2551:
2521:
2520:
2516:
2472:
2471:
2464:
2434:
2433:
2424:
2394:
2393:
2389:
2359:
2358:
2351:
2317:
2316:
2309:
2287:
2286:
2282:
2251:(6800): 81–86.
2238:
2237:
2224:
2193:
2192:
2175:
2142:(2): e1000313.
2129:
2128:
2099:
2063:
2062:
2058:
2004:
2003:
1999:
1962:(6850): 904–7.
1953:
1952:
1948:
1902:
1901:
1897:
1847:
1846:
1842:
1821:(6–7): 488–96.
1808:
1807:
1800:
1762:
1761:
1757:
1719:
1718:
1714:
1668:
1667:
1663:
1596:
1595:
1578:
1571:
1556:
1555:
1551:
1529:
1528:
1524:
1488:
1487:
1480:
1430:
1429:
1422:
1413:
1402:
1397:
1393:
1360:(2): e1000313.
1347:
1346:
1337:
1332:
1325:
1312:
1310:
1302:
1301:
1297:
1278:
1277:
1273:
1261:
1260:
1256:
1238:
1237:
1233:
1228:
1200:
1169:
1100:
1027:
999:
931:Medicago sativa
918:
874:
821:
810:
804:
801:
793:help improve it
790:
781:
777:
770:
513:
477:species complex
465:cryptic species
454:scientific name
381:
377:
376:
304:parthenogenesis
297:sexual maturity
285:
196:A. pisum ononis
175:
169:
163:
150:
48:
17:
12:
11:
5:
3288:
3286:
3278:
3277:
3272:
3267:
3262:
3257:
3252:
3247:
3242:
3232:
3231:
3225:
3224:
3222:
3221:
3211:
3198:
3188:
3175:
3162:
3149:
3136:
3123:
3110:
3097:
3088:Fauna Europaea
3084:
3071:
3058:
3045:
3032:
3019:
3006:
2993:
2978:
2962:
2960:
2954:
2953:
2948:
2940:
2939:
2889:
2848:
2791:
2761:
2723:
2688:
2612:
2549:
2530:(1): 203–210.
2514:
2462:
2443:(4): 747–754.
2422:
2387:
2349:
2307:
2280:
2222:
2173:
2097:
2056:
2017:(29): 9965–9.
1997:
1946:
1895:
1840:
1798:
1771:(5666): 1989.
1755:
1728:(5755): 1781.
1712:
1661:
1620:BioMed Central
1611:Genome Biology
1576:
1569:
1549:
1522:
1478:
1420:
1400:
1391:
1335:
1323:
1295:
1271:
1254:
1230:
1229:
1227:
1224:
1199:
1196:
1184:bacterium and
1168:
1165:
1099:
1096:
1026:
1023:
998:
995:
917:
914:
873:
870:
823:
822:
784:
782:
775:
769:
763:
762:
761:
748:
730:
693:
681:
665:
552:hemimetabolous
512:
511:Model organism
509:
284:
281:
265:model organism
222:
221:
220:
219:
210:
201:
193:
188:A. pisum pisum
182:
181:
177:
176:
170:
159:
158:
152:
151:
144:
142:
138:
137:
130:
126:
125:
120:
116:
115:
113:Sternorrhyncha
110:
106:
105:
100:
96:
95:
90:
86:
85:
80:
76:
75:
70:
66:
65:
60:
56:
55:
42:
41:
37:
36:
28:
27:
15:
13:
10:
9:
6:
4:
3:
2:
3287:
3276:
3273:
3271:
3268:
3266:
3263:
3261:
3258:
3256:
3253:
3251:
3248:
3246:
3243:
3241:
3238:
3237:
3235:
3218:
3212:
3208:
3203:
3199:
3195:
3189:
3185:
3180:
3176:
3172:
3167:
3163:
3159:
3154:
3150:
3146:
3141:
3137:
3133:
3128:
3124:
3120:
3115:
3111:
3107:
3102:
3098:
3094:
3089:
3085:
3081:
3076:
3072:
3068:
3063:
3059:
3055:
3050:
3046:
3042:
3037:
3033:
3029:
3024:
3020:
3016:
3011:
3007:
3003:
2998:
2994:
2989:
2983:
2979:
2974:
2968:
2964:
2963:
2961:
2959:
2955:
2951:
2946:
2935:
2931:
2927:
2923:
2919:
2915:
2911:
2907:
2900:
2898:
2896:
2894:
2890:
2885:
2881:
2876:
2871:
2867:
2863:
2862:Mol Biol Evol
2859:
2852:
2849:
2844:
2840:
2836:
2832:
2827:
2822:
2818:
2814:
2810:
2806:
2802:
2795:
2792:
2787:
2783:
2779:
2775:
2768:
2766:
2762:
2757:
2753:
2749:
2745:
2741:
2737:
2730:
2728:
2724:
2719:
2715:
2711:
2707:
2703:
2699:
2692:
2689:
2684:
2680:
2675:
2670:
2666:
2662:
2658:
2654:
2650:
2643:
2641:
2639:
2637:
2635:
2633:
2631:
2629:
2627:
2625:
2623:
2621:
2619:
2617:
2613:
2608:
2604:
2600:
2596:
2592:
2588:
2584:
2580:
2572:
2570:
2568:
2566:
2564:
2562:
2560:
2558:
2556:
2554:
2550:
2545:
2541:
2537:
2533:
2529:
2525:
2518:
2515:
2510:
2506:
2501:
2496:
2492:
2488:
2484:
2480:
2476:
2469:
2467:
2463:
2458:
2454:
2450:
2446:
2442:
2438:
2431:
2429:
2427:
2423:
2418:
2414:
2410:
2406:
2402:
2398:
2391:
2388:
2383:
2379:
2375:
2371:
2367:
2363:
2356:
2354:
2350:
2345:
2341:
2337:
2333:
2329:
2325:
2321:
2314:
2312:
2308:
2303:
2299:
2295:
2291:
2284:
2281:
2276:
2272:
2267:
2262:
2258:
2254:
2250:
2246:
2242:
2235:
2233:
2231:
2229:
2227:
2223:
2218:
2214:
2210:
2206:
2202:
2198:
2190:
2188:
2186:
2184:
2182:
2180:
2178:
2174:
2169:
2165:
2160:
2155:
2150:
2145:
2141:
2137:
2133:
2126:
2124:
2122:
2120:
2118:
2116:
2114:
2112:
2110:
2108:
2106:
2104:
2102:
2098:
2092:
2087:
2083:
2079:
2075:
2071:
2067:
2060:
2057:
2052:
2048:
2043:
2038:
2033:
2028:
2024:
2020:
2016:
2012:
2008:
2001:
1998:
1993:
1989:
1985:
1981:
1977:
1973:
1969:
1965:
1961:
1957:
1950:
1947:
1942:
1938:
1934:
1930:
1926:
1922:
1918:
1914:
1910:
1906:
1899:
1896:
1891:
1887:
1883:
1879:
1875:
1871:
1867:
1863:
1859:
1855:
1851:
1844:
1841:
1836:
1832:
1828:
1824:
1820:
1816:
1812:
1805:
1803:
1799:
1794:
1790:
1786:
1782:
1778:
1774:
1770:
1766:
1759:
1756:
1751:
1747:
1743:
1739:
1735:
1731:
1727:
1723:
1716:
1713:
1708:
1704:
1700:
1696:
1692:
1688:
1684:
1680:
1676:
1672:
1665:
1662:
1657:
1653:
1649:
1645:
1640:
1635:
1630:
1625:
1621:
1617:
1613:
1612:
1607:
1605:
1599:
1593:
1591:
1589:
1587:
1585:
1583:
1581:
1577:
1572:
1566:
1562:
1561:
1553:
1550:
1545:
1541:
1537:
1533:
1526:
1523:
1518:
1514:
1509:
1504:
1500:
1496:
1492:
1485:
1483:
1479:
1474:
1470:
1465:
1460:
1455:
1450:
1446:
1442:
1438:
1436:
1427:
1425:
1421:
1417:
1416:Acyrthosiphon
1411:
1409:
1407:
1405:
1401:
1395:
1392:
1387:
1383:
1378:
1373:
1368:
1363:
1359:
1355:
1351:
1344:
1342:
1340:
1336:
1330:
1328:
1324:
1320:
1309:
1305:
1299:
1296:
1292:
1288:
1284:
1283:
1275:
1272:
1267:
1266:
1258:
1255:
1250:
1246:
1242:
1235:
1232:
1225:
1223:
1221:
1217:
1216:
1213:
1208:
1204:
1197:
1195:
1193:
1191:
1187:
1183:
1178:
1174:
1167:Immune system
1166:
1164:
1162:
1158:
1153:
1149:
1145:
1141:
1136:
1134:
1130:
1126:
1122:
1118:
1114:
1109:
1105:
1097:
1095:
1093:
1089:
1085:
1080:
1077:
1072:
1067:
1064:
1060:
1056:
1053:
1049:
1044:
1040:
1035:
1031:
1024:
1022:
1020:
1016:
1011:
1007:
1003:
996:
994:
992:
988:
984:
980:
976:
972:
968:
964:
959:
955:
951:
950:
945:
944:
939:
938:
933:
932:
927:
923:
915:
913:
911:
907:
903:
899:
895:
891:
887:
883:
879:
871:
869:
867:
863:
859:
855:
851:
847:
843:
839:
835:
834:
829:
819:
816:
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785:This section
783:
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739:
738:gene families
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496:North America
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463:described as
462:
458:
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411:
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401:that feed on
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327:overwintering
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160:
157:
156:Binomial name
153:
149:
148:
147:A. pisum
143:
140:
139:
136:
135:
134:Acyrthosiphon
131:
128:
127:
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121:
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29:
26:
22:
19:
3240:Macrosiphini
2957:
2917:
2913:
2909:
2905:
2865:
2861:
2851:
2808:
2804:
2794:
2777:
2773:
2742:(1): 41–46.
2739:
2735:
2701:
2697:
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2656:
2652:
2590:
2586:
2582:
2578:
2527:
2523:
2517:
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2436:
2403:(1): 31–38.
2400:
2396:
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2327:
2323:
2319:
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2248:
2244:
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2196:
2139:
2136:PLOS Biology
2135:
2073:
2069:
2059:
2014:
2010:
2000:
1959:
1955:
1949:
1908:
1904:
1898:
1860:(5859): 58.
1857:
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1814:
1768:
1764:
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1725:
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1552:
1535:
1531:
1525:
1501:(3): 192–9.
1498:
1494:
1444:
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1357:
1354:PLOS Biology
1353:
1318:
1311:. Retrieved
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953:
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937:Pisum sativa
935:
929:
925:
921:
919:
909:
905:
901:
897:
893:
889:
885:
881:
877:
875:
865:
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849:
846:holosymbiont
845:
841:
837:
832:
831:
827:
826:
811:
802:
786:
766:
750:
742:
732:
710:
697:Polymorphism
695:
683:
667:
656:
650:
640:
634:
620:horizontally
609:
603:
597:
593:
589:
579:
577:endosymbiont
572:
558:
527:
526:
504:
491:
481:
456:
451:
446:
431:insecticides
418:
413:
407:
393:through its
372:
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308:
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232:
227:
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225:
213:
204:
195:
187:
164:
162:
146:
145:
133:
24:
18:
3166:NatureServe
3127:iNaturalist
2982:Wikispecies
2920:: 241–248.
2780:: 115–135.
2593:: 249–258.
2076:: 119–130.
1538:(2): 3–16.
1220:biocontrols
1066:bacterium.
934:(alfalfa),
715:carotenoids
669:Polyphenism
586:amino acids
548:chromosomes
435:parasitoids
367:populations
323:Inseminated
180:Subspecies
3234:Categories
1447:(39): 39.
1226:References
1215:neoaphidis
949:Vicia faba
852:bacteria,
753:speciation
744:Drosophila
701:phenotypic
661:neoaphidis
616:maternally
539:and 34000
484:Palearctic
469:subspecies
391:phloem sap
359:hereditary
347:Inbreeding
341:diapausing
315:parasitism
301:viviparous
257:broad bean
208:Koch, 1855
199:Koch, 1855
109:Suborder:
83:Arthropoda
2368:: 55–94.
2330:: 17–37.
1707:206529034
1088:Buchnera.
1059:A. pisum.
991:A. pisum.
805:June 2014
569:Aphididae
567:- As all
560:Symbiosis
541:predicted
537:megabases
439:cultivars
421:can be a
403:honeydews
337:asexually
319:oviparous
311:predation
277:annotated
273:sequenced
271:has been
237:Aphididae
233:pea aphid
141:Species:
123:Aphididae
103:Hemiptera
69:Kingdom:
63:Eukaryota
3171:2.895849
3145:11057434
3036:BugGuide
2973:Q1348560
2967:Wikidata
2934:20482654
2884:21613235
2843:20882842
2835:21692233
2805:Mol Ecol
2683:21390549
2607:20482655
2509:19129128
2457:16449374
2344:15012383
2320:Buchnera
2275:10993077
2217:19909925
2168:20186266
2051:18621720
1984:11528477
1941:14785276
1933:20431015
1890:34320106
1882:18174433
1835:20541160
1793:46139967
1785:15044797
1750:32543175
1742:16357252
1699:21097935
1648:20178569
1517:16823401
1495:Heredity
1473:22954315
1386:20186266
1313:19 March
1251:: 1–212.
1203:A. pisum
1190:A. pisum
1186:A. pisum
1182:Buchnera
1177:A. pisum
1173:A. pisum
1161:Buchnera
1157:A. pisum
1152:Buchnera
1148:A. pisum
1144:A. pisum
1140:Buchnera
1133:Buchnera
1129:A. pisum
1125:A. pisum
1121:Buchnera
1117:A. pisum
1113:Buchnera
1108:Buchnera
1104:A. pisum
1092:Buchnera
1084:A. pisum
1076:A. pisum
1071:A. pisum
1063:Buchnera
1055:Buchnera
1052:E. coli.
1048:Buchnera
1043:Buchnera
1039:A. pisum
1034:Buchnera
1030:A. pisum
1019:Buchnera
1015:Buchnera
1010:Buchnera
1006:A. pisum
1002:Buchnera
987:Buchnera
983:Buchnera
979:A. pisum
975:Buchnera
971:A. pisum
967:Buchnera
963:A. pisum
954:A. pisum
926:A. pisum
910:Buchnera
902:Buchnera
898:Buchnera
894:A. pisum
890:Buchnera
886:Buchnera
882:Buchnera
878:A. pisum
866:Buchnera
862:A. pisum
858:Buchnera
854:A. pisum
850:Buchnera
842:Buchnera
838:A. pisum
828:A. pisum
723:carotene
719:torulene
711:A. pisum
674:genotype
594:A. pisum
590:Buchnera
573:A. pisum
564:bacteria
546:in 2n=8
528:A. pisum
492:A. pisum
461:biotypes
457:A. pisum
447:A. pisum
419:A. pisum
414:A. pisum
352:Several
241:Fabaceae
119:Family:
79:Phylum:
73:Animalia
59:Domain:
3119:2077503
2813:Bibcode
2744:Bibcode
2706:Bibcode
2674:3064905
2532:Bibcode
2500:2664372
2405:Bibcode
2382:8561471
2253:Bibcode
2159:2826372
2078:Bibcode
2042:2481341
2019:Bibcode
1992:4325260
1964:Bibcode
1913:Bibcode
1905:Science
1862:Bibcode
1854:Science
1765:Science
1722:Science
1679:Bibcode
1671:Science
1656:5249535
1639:2872881
1622:: R21.
1464:3471804
1377:2826372
1212:Pandora
969:allows
958:de novo
940:(pea),
922:A.pisum
791:Please
678:genetic
658:Pandora
624:abiotic
521:alfalfa
395:stylets
380:⁄
253:alfalfa
129:Genus:
99:Order:
93:Insecta
89:Class:
3207:814484
3191:NZOR:
3158:200572
3132:132882
3080:ACYRON
3067:588004
3041:356333
2932:
2882:
2841:
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2681:
2671:
2605:
2507:
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1990:
1982:
1956:Nature
1939:
1931:
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1567:
1515:
1471:
1461:
1384:
1374:
1289:–154.
649:, and
631:stress
628:biotic
423:vector
410:genera
363:morphs
354:morphs
293:moults
289:nymphs
269:genome
255:, and
249:clover
191:(type)
174:, 1776
172:Harris
3214:PPE:
3140:IRMNG
3093:54351
3028:14527
2839:S2CID
1988:S2CID
1937:S2CID
1886:S2CID
1789:S2CID
1746:S2CID
1703:S2CID
1652:S2CID
1618:(2).
1308:AphID
562:with
544:genes
473:races
441:. No
427:viral
261:aphid
3184:7029
3179:NCBI
3153:ITIS
3114:GBIF
3075:EPPO
3054:9WK9
3023:BOLD
2930:PMID
2880:PMID
2831:PMID
2679:PMID
2603:PMID
2505:PMID
2453:PMID
2378:PMID
2340:PMID
2271:PMID
2213:PMID
2164:PMID
2047:PMID
1980:PMID
1929:PMID
1878:PMID
1831:PMID
1781:PMID
1738:PMID
1695:PMID
1644:PMID
1565:ISBN
1513:PMID
1469:PMID
1382:PMID
1315:2013
1159:and
1150:and
1138:The
1131:and
1106:and
1086:and
1069:The
1061:The
1041:and
1032:and
892:and
880:and
876:The
864:and
707:Loci
639:and
626:and
618:and
399:ants
387:pods
330:eggs
313:and
275:and
3062:EoL
3049:CoL
3010:AFD
2997:ADW
2922:doi
2912:".
2870:doi
2821:doi
2782:doi
2752:doi
2714:doi
2669:PMC
2661:doi
2595:doi
2585:".
2540:doi
2495:PMC
2487:doi
2483:276
2445:doi
2413:doi
2370:doi
2332:doi
2322:".
2298:doi
2294:253
2261:doi
2249:407
2205:doi
2201:332
2154:PMC
2144:doi
2086:doi
2037:PMC
2027:doi
2015:105
1972:doi
1960:412
1921:doi
1909:328
1870:doi
1858:319
1823:doi
1819:333
1773:doi
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1730:doi
1726:310
1687:doi
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1634:PMC
1624:doi
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1503:doi
1459:PMC
1449:doi
1372:PMC
1362:doi
1287:138
795:to
507:).
471:or
425:of
245:pea
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