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Acyrthosiphon pisum
Acyrthosiphon pisum , comúnmente conocido como el áfido del
[1] [2] Áfido de guisante
guisante (y coloquialmente conocido como el delfín verde, piojo
del guisante, y piojo del trébol [3] ), es un insecto succionador de savia en
lafamilia Aphididae . Se alimenta de varias especies de leguminosas
(familia de plantas Fabaceae ) en todo el mundo, incluidos los cultivos
de forraje, como el guisante , el trébol , la alfalfa y el haba , [4] y se
encuentra entre lasespecies de áfidos de mayor importancia agronómica.
[5] El áfido del guisante es un organismo modelo para el estudio biológico
cuyoel genoma ha sido secuenciado y anotado . [6]

Contenido
Adulto partenogenético, áfido de
Generalidades y ciclo de vida guisante y progenie alimentándose en
organismo modelo
una planta de guisante
Relación endosimbiótica con Buchnera aphidicola
Evolución de la relación endosimbiótica clasificación cientifica
simbiosis nutricional Reino: Animalia
Estructura Holosymbiont
Secuenciación del genoma Filo: Arthropoda
Complementariedad metabólica
Clase: Insecta
Sistema inmune
referencias
Orden: Homoptera
Suborden: Sternorrhyncha
Superfamilia: Aphidoidea
Generalidades y ciclo de vida Familia: Aphididae
En otoño, los áfidos de las arvejas ponen huevos fertilizados durante el
Género: Acyrthosiphon
invierno que nacen en la primavera siguiente. Las ninfas que nacen de
Mordvilko de 1914
estos huevos son todas hembras, que se someten a cuatro mudas antes
de alcanzar la madurez sexual . Luego comenzarán a reproducirse por Especies: A. pisum
partenogénesis vivípara , como la mayoría de los áfidos. Cada hembra
Nombre binomial
adulta da a luz de cuatro a 12 ninfas hembras por día, alrededor de cien
en su vida. Estos se convierten en hembras maduras en Acyrthosiphon pisum
aproximadamente siete a diez días. La vida útil de un adulto es de Harris , 1776
aproximadamente 30 días.
Subespecie
Las densidades de población están en su punto más alto a principios del
A. pisum pisum (tipo)
verano, luego disminuyen a través de la depredación y el parasitismo .
En otoño, el alargamiento de la noche desencadena la producción de una A. pisum ononis Koch, 1855
sola generación de individuos sexuales (varones y hembras ovíparas ) ? A. pisum spartii Koch, 1855
por las mismas hembras progenitoras partenogenéticas. Las hembras
? A. pisum destructor Johnson, 1900
sexuales inseminadas pondrán huevos para invernar , de los cuales
surgirán nuevas hembras partenogenéticas a comienzos de la primavera.
Cuando la colonia comienza a estar abarrotada, se producen algunas hembras aladas. Estos se dispersan para infestar

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otras plantas, donde continúan reproduciéndose asexualmente. Cuando las temperaturas se vuelven más frías y la
duración del día es más corta, aparecen las hembras y los machos con alas sexuales. Estas compañeras, las hembras
ponen huevos para diapaso y el ciclo de vida comienza de nuevo. [7] Los áfidos de los guisantes pueden completar todo
su ciclo reproductivo sin cambiar la planta huésped. [8]
Existen varios morfos en los áfidos de los guisantes. Además de las diferencias entre los morfos sexuales y
partenogenéticos, existen morfos alados y sin alas. La sobrepoblación y la mala calidad de los alimentos pueden
desencadenar el desarrollo de individuos alados en las generaciones posteriores. [9] Los áfidos con alas pueden
colonizar otras plantas hospederas. Los áfidos de guisante también muestran variaciones de color de cuerpo
hereditario de verde o rojo / rosa. Los morfos verdesson generalmente más frecuentes en poblaciones naturales . [8]

Acyrthosiphon pisum es un áfido bastante grande cuyo cuerpo puede alcanzar 4 mm en adultos. [8] Por lo general, se
alimenta de los lados inferiores de las hojas, los brotes y las vainas de las leguminosas, ingiriendo savia de floema a
través de sus estiletes . A diferencia de muchas especies de áfidos, los áfidos de los guisantes no tienden a formar
colonias densas donde las personas se quedarían donde nacieron durante toda su vida. No se sabe que los áfidos de los
guisantes sean cultivados por hormigas que se alimentan de honeydews .

Se sabe que más de 20 géneros de leguminosas albergan áfidos de guisante, aunque el rango de hospedadores
completo permanece indeterminado. En cultivos tales como guisantes y alfalfa, A. pisum se considera entre las
especies de áfidos o de mayor importancia agronómica. [5] Los rendimientos pueden verse afectados por la ingesta de
savia que debilita directamente a las plantas, aunque los áfidos de los guisantes rara vez alcanzan densidades que
pueden reducir significativamente la producción de cultivos. Sin embargo, como muchas especies de áfidos, A. pisum
puede ser un vector de enfermedades virales para las plantas que visita. La protección contra los áfidos de los
guisantes incluye el uso de insecticidas químicos , depredadores naturales y parasitoides , y la selección de
resistentescultivares . No se documenta resistencia a insecticidas en A. pisum , a diferencia de muchas plagas de
áfidos.

Los áfidos de los guisantes, aunque designados colectivamente por el único nombre científico A. pisum , abarcan
varios biotipos descritos como especies crípticas , subespecies o razas , que están especializadas en diferentes especies
hospedadoras. Por lo tanto, el áfido del guisante se describe con mayor precisión como un complejo de especies . [8]

Se cree que el áfido del guisante es de origen paleártico , pero ahora se encuentra comúnmente en todo el mundo en
climas templados . La propagación de A. pisum probablemente se debió a la introducción de algunas de sus plantas
hospedantes para la agricultura. Tal introducción probablemente ocurrió en América del Norte durante la década de
1870, [10] y en 1900 se había convertido en una especie de plaga grave en los estados del Atlántico medio. En la década
de 1950, se extendió por todos los Estados Unidos y Canadá. Su rango de hospederos en América del Norte es muy
similar al del áfido azul de la alfalfa ( Acyrthosiphon kondoi ). [11]

Organismo modelo
A. pisum se considera como la especie de áfido modelo. Su ciclo reproductivo, incluida la fase sexual y el hibernación
de los huevos, se puede completar fácilmente en las plantas hospedadoras en condiciones de laboratorio, y el tamaño
relativamente grande de las personas facilita los estudios fisiológicos. En 2010, el International Aphid Genomics
Consortium publicó una secuencia preliminar anotada del genoma del áfido del guisante [6] compuesta de
aproximadamente 525 megabases y 34000 genes pronosticados en 2n = 8 cromosomas . Esto constituye el primer
genoma de un hemimetabolousinsecto para haber sido publicado. El genoma del áfido del guisante y otras
características son el foco de los estudios que cubren las siguientes áreas:

Simbiosis con bacterias : como todos los aphididae , A. pisum alberga el endosimbionte primario Buchnera
aphidicola , que proporciona aminoácidos esencialesy es necesario para la reproducción de los áfidos. Buchnera
se transmite de madres a crías, y ha coevolucionado con áfidos durante decenas de millones de años. A. pisum
también alberga una gama de simbiontes bacterianos facultativos que pueden transmitirse por vía materna y
horizontal , y que afectan los rasgos ecológicamente importantes en los áfidos, como el color del cuerpo, [12] la
resistencia a los abióticos y estrés biótico , [13] y nutrición. [14]
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Polifenismo (la producción de varios morfos discretos por el mismo

genotipo ) - Los estudios sobre áfidos de guisante han ayudado a
establecer loscomponentesambientales y genéticos que controlan la
producción demorfossexuales [15] y alados, [9] entre otras
características.
Reproducción asexual : los linajes del áfido del guisante incluyen la
partenogénesis en sus ciclos de vida, y algunos incluso han perdido la
fase sexual. Los áfidos de guisante son modelos para descifrar el
origen y las consecuencias de la reproducción asexual, [15] [16] una
cuestión importante en la biología evolutiva .
Polymorphism and physiology explaining phenotypic variations
in aphids - Loci 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. A. pisum is notable for being the only animal organism
so-far identified that has the ability to synthesize a carotenoid. Plants, Adulto, aspidán de guisante
fungi, and microorganisms can synthesize carotenoids, but torulene partenogenético en alfalfa - este
(3',4'-didehydro-β,γ-carotene, specifically a hydrocarbon carotene) morfo rojo muestra las marcas
made by pea aphids, is the only carotenoid known to be synthesized
rojizas / oscuras debido a los
by an organism in the animal kingdom. Torulene imparts natural, red-
colored patches to some aphids, which possibly aid in their carotenoides que producen algunas
camouflage and escape from predation by wasps. The aphids have personas.
gained the ability to synthesize torulene by horizontal gene transfer of
a number of genes for carotenoid synthesis, apparently from fungi.[17]
Gene duplication and expansion of gene families - The pea aphid genome presents high levels of gene
duplication compared to other insect genomes, such as Drosophila, with the notable expansion of some gene
families.[6]
Interaction with host plants and speciation - 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,[18] molecular and physiological
mechanisms that are involved in host nutrition and virulence.[19] Genetic, molecular and physiological studies
have also evidenced specialization to different host species as a motor of ecological speciation between pea
aphid biotypes.[20]

Endosymbiotic Relationship with Buchnera aphidicola

A. pisum participates in an obligate endosymbiotic relationship with the bacteria Buchnera aphidicola. A. pisum is
the host and Buchnera is the primary endosymbiont. Together they form the holosymbiont.[21] This is an obligate,
symbiotic relationship and both partners are completely dependent on each other.[21][22] When treated with antibiotics
to remove the Buchnera bacteria, A. pisum growth and reproduction are interrupted or reduced. Buchnera lacks genes
required for living independent of a host and is unculturable outside of the aphid host.[23] The A. pisum and Buchnera
holosymbiont is one of the most well studied symbiotic relationships both genetically and experimentally.

Evolution of the Endosymbiotic Relationship

The A. pisum and Buchnera endosymbiotic relationship is likely to have evolved 160-280 million years ago.
Phylogenetic analysis shows that Buchnera is a monophyletic group and that the phylogenies of Buchnera and A.
pisum coincide. Therefore, there was likely one original Buchnera infection of the common ancestor of aphids and co-
speciation of the holosymbiont has occurred since then.[24] Buchnera is related to Enterobacteriaceae including
Escheriachia coli [22] and it is likely that Buchnera evolved from a bacterium that originally occupied the gut of the
aphid common ancestor.[25]

Nutritional symbiosis
Like other insects of the order Hemiptera, A.pisum utilizes an endosymbiotic bacterium to overcome the nutritional
deficiencies of phloem sap.[21][26] A. pisum feeds on phloem sap of host plants including Medicago sativa (alfalfa),
Pisum sativa (pea), Trifolium pretense (red clover), and Vicia faba (broad bean). The phloem saps of these plants are
nutritionally rich in carbohydrates but poor in terms of nitrogen.[27][28][29][30] The ratio of essential amino acids to

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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.[28]
Animals, including A. pisum, can produce nonessential amino acids de novo 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, A. pisum is unable
to synthesize arginine due to missing urea cycle genes.[21][31][32] The endosymbiotic relationship with Buchnera allows
A. pisum to overcome this lack of essential amino acids in the phloem sap [22][28][29][32][33] When provided with
nonessential amino acids, Buchnera converts nonessential amino acids into essential amino acids to be returned to A.
pisum.[26][34] This nutritional provisioning has been examined genomically (metabolic complementary, discussed
below) and experimentally. Isolated bacteriocytes containing Buchnera have been shown to actively take up 14C
labeled glutamine (a nonessential amino acid) where it is then converted into glutamic acid.[34] This glutamic acid is
then taken up by the individual Buchnera 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 A. pisum. Mutual nutrient
provisioning is likely the main reason for the persistence of this symbiosis.[32]

Holosymbiont Structure
Buchnera are housed in specialized, aphid-derived cells located in the hemocoel of the A. pisum body cavity.[21][22]
Each Buchnera 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 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.[22][31][35]
Bacteriocytes are located near the ovariole cluster and Buchnera cells are vertically transferred from the mother’s
ovaries through transovarial transmission.[22][25] The Buchnera cells are transferred to eggs during oogenesis or to the
developing embryos during embryogenesis.[21]

Genome Sequencing
A. pisum and Buchnera were the first insect-endosymbiont pair to have the genomes of both partners sequenced.[36]
This has provided researchers with a great deal of information about the evolutionary and molecular interactions of
this endosymbiosis.[31] The A. pisum and Buchnera 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 Buchnera 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,
E. coli.[23][31] Buchnera has lost genes that would allow it to live outside the host but maintains genes essential for the
nutrition of A. pisum.[21][22][23][31][32] The Buchnera genome is missing genes required for surface membrane
construction such as lipopolysaccharides and phosopholipids 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 bacterium.[23][32]

The A. pisum 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.[21][32] These orphan genes and gene duplications are likely
associated with the “metabolic, structural and developmental” components of the endosymbiotic relationship.[32] A.
pisum specific gene duplications of amino acid transporters highly expressed in bacteriocytes have been observed.[37]
These duplications are likely associated with the genetic establishment and maintenance of the endosymbiotic
relationship.

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No lateral gene transfer has been detected between A. pisum and Buchnera. It was previously believed that lateral
gene transfer was responsible for the severe gene reduction in the Buchnera genome but sequencing has shown that
this has not occurred.[23][32]

Metabolic complementarity
Individually, the metabolic pathways of A. pisum and Buchnera 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.[21][31][32][38] The ancestral partners of this symbiosis are likely to have had
complete metabolic pathways, however pressure to maintain these pathway genes was reduced due to redundancy as a
result of the presence of the other partner’s genome.[38] Unlike other related insects, the A. pisum genome is missing
genes necessary for the urea cycle.[21][31][35] the purine salvage pathway,[38] and other genes that code enzymes
necessary for the biosynthesis of molecules.[31][32] These missing reaction intermediates are likely provided by genes
within the Buchnera genome. For example, A. pisum 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 Buchnera genome contains the necessary genes to encode the reaction intermediates missing
from the A. pisum genome. Through this complementation, the nucleotide requirements of both organisms are
fulfilled: the purine salvage pathway is completed for A. pisum and Buchnera receives necessary guanosine.[38]

The Buchnera 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 A. pisum genome on the other hand, contains 66 amino acid
biosynthesis genes and 93 amino acid degradation genes.[21][31] Both A. pisum and Buchnera contribute to the
metabolic pathways of amino acid biosynthesis.[21][31][32] 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 A. pisum and Buchnera.[32]

Immune system
Genome sequencing of A. pisum shows that the genome lacks expected genes essential to immune response
pathways.[39] The A. pisum 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 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 Buchnera bacterium and A.
pisum.[21][32] Also, phloem sap is a diet with reduced amounts of microbes which may have lower the evolutionary
pressure of A. pisum to maintain the immune response pathway genes.[32]

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