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vely on sap of phloem vessels in plants as do many other hemipterans such as scale insects and cicadas once a phloem vessel is punctured the sap which is under pressure is forced into the aphid s food canal occasionally aphids also ingest xylem sap which is a more dilute diet than phloem sap as the concentrations of sugars and amino acids are 1 of those in the phloem 36 37 xylem sap is under negative hydrostatic pressure and requires active sucking suggesting an important role in aphid physiology 38 as xylem sap ingestion has been observed following a dehydration period aphids are thought to consume xylem sap to replenish their water balance the consumption of the dilute sap of xylem permitting aphids to rehydrate 39 however recent data showed aphids consume more xylem sap than expected and they notably do so when they are not dehydrated and when their fecundity decreases this suggests aphids and potentially all the phloem sap feeding species of the order hemiptera consume xylem sap for reasons other than replenishing water balance 40 although aphids passively take in phloem sap which is under pressure they can also draw fluid at negative or atmospheric pressure using the cibarial pharyngeal pump mechanism present in their head 41 xylem sap consumption may be related to osmoregulation 40 high osmotic pressure in the stomach caused by high sucrose concentration can lead to water transfer from the hemolymph to the stomach thus resulting in hyperosmotic stress and eventually to the death of the insect aphids avoid this fate by osmoregulating through several processes sucrose concentration is directly reduced by assimilating sucrose toward metabolism and by synthesizing oligosaccharides from several sucrose molecules thus reducing the solute concentration and consequently the osmotic pressure 42 43 oligosaccharides are then excreted through honeydew explaining its high sugar concentrations which can then be used by other animals such as ants furthermore water is transferred from the hindgut where osmotic pressure has already been reduced to the stomach to dilute stomach content 44 eventually aphids consume xylem sap to dilute the stomach osmotic pressure 40 all these processes function synergetically and enable aphids to feed on high sucrose concentration plant sap as well as to adapt to varying sucrose concentrations 40 plant sap is an unbalanced diet for aphids as it lacks essential amino acids which aphids like all animals cannot synthesise and possesses a high osmotic pressure due to its high sucrose concentration 37 45 essential amino acids are provided to aphids by bacterial endosymbionts harboured in special cells bacteriocytes 46 these symbionts recycle glutamate a metabolic waste of their host into essential amino acids 47 48 carotenoids and photoheterotrophy edit some species of aphids have acquired the ability to synthesise red carotenoids by horizontal gene transfer from fungi 49 they are the only animals other than two spotted spider mites and the oriental hornet with this capability 50 using their carotenoids aphids may well be able to absorb solar energy and convert it to a form that their cells can use atp this is the only known example of photoheterotrophy in animals the carotene pigments in aphids form a layer close to the surface of the cuticle ideally placed to absorb sunlight the excited carotenoids seem to reduce nad to nadh which is oxidized in the mitochondria for energy 51 reproduction edit soybean aphid alternates between hosts and between asexual and sexual reproduction 52 winged adult alate milkweed aphids some scenes repeated at one twentieth speed recorded at 60 fps the simplest reproductive strategy is for an aphid to have a single host all year round on this it may alternate between sexual and asexual generations holocyclic or alternatively all young may be produced by parthenogenesis eggs never being laid anholocyclic some species can have both holocyclic and anholocyclic populations under different circumstances but no known aphid species reproduce solely by sexual means 53 the alternation of sexual and asexual generations may have evolved repeatedly 54 however aphid reproduction is often more complex than this and involves migration between different host plants in about 10 of species there is an alternation between woody primary hosts on which the aphids overwinter and herbaceous secondary host plants where they reproduce abundantly in the summer 21 53 a few species can produce a soldier caste other species show extensive polyphenism under different environmental conditions and some can control the sex ratio of their offspring depending on external factors 55 when a typical sophisticated reproductive strategy is used only females are present in the population at the beginning of the seasonal cycle although a few species of aphids have been found to have both male and female sexes at this time the overwintering eggs that hatch in the spring result in females called fundatrices stem mothers reproduction typically does not involve males parthenogenesis and results in a live birth viviparity 56 the live young are produced by pseudoplacental viviparity which is the development of eggs deficient in the yolk the embryos fed by a tissue acting as a placenta the young emerge from the mother soon after hatching 57 eggs are parthenogenetically produced without meiosis 58 56 and the offspring are clonal to their mother so they are all female thelytoky 12 57 the embryos develop within the mothers ovarioles which then give birth to live already hatched first instar female nymphs as the eggs begin to develop immediately after ovulation an adult female can house developing female nymphs which already have parthenogenetically developing embryos inside them i e they are born pregnant this telescoping of generations enables aphids to increase in number with great rapidity the offspring resemble their parent in every way except size thus a female s diet can affect the body size and birth rate of more than two generations daughters and granddaughters 12 59 60 aphid giving birth to live young populations are often entirely female this process repeats itself throughout the summer producing multiple generations that typically live 20 to 40 days for example some species of cabbage aphids like brevicoryne brassicae can produce up to 41 generations of females in a season thus one female hatched in spring can theoretically produce billions of descendants were they all to survive 61 in autumn aphids reproduce sexually and lay eggs environmental factors such as a change in photoperiod and temperature or perhaps a lower food quantity or quality causes females to parthenogenetically produce sexual females and males 58 the males are genetically identical to their mothers except that with the aphids x0 sex determination system they have one fewer sex chromosome 58 these sexual aphids may lack wings or even mouthparts 21 sexual females and males mate and females lay eggs that develop outside the mother the eggs survive the winter and hatch into winged alate or wingless females the following spring this occurs in for example the life cycle of the rose aphid macrosiphum rosae which may be considered typical of the family however in warm environments such as in the tropics or a greenhouse aphids may go on reproducing asexually for many years 29 the life stages of the green apple aphid aphis pomi drawing by robert evans snodgrass 1930 aphids reproducing asexually by parthenogenesis can have genetically identical winged and non winged female progeny control is complex some aphids alternate during their life cycles between genetic control polymorphism and environmental control polyphenism of production of winged or wingless forms 62 winged progeny tend to be produced more abundantly under unfavorable or stressful conditions some species produce winged progeny in response to low food quality or quantity e g when a host plant is starting to senesce 63 the winged females migrate to start new colonies on a new host plant for example the apple aphid aphis pomi after producing many generations of wingless females gives rise to winged forms that fly to other branches or trees of its typical food plant 64 aphids that are attacked by ladybugs lacewings parasitoid wasps or other predators can change the dynamics of their progeny production when aphids are attacked by these predators alarm pheromones in particular beta farnesene are released from the cornicles these alarm pheromones cause several behavioral modifications that depending on the aphid species can include walking away and dropping off the host plant additionally alarm pheromone perception can induce the aphids to produce winged progeny that can leave the host plant in search of a safer feeding site 65 viral infections which can be extremely harmful to aphids can also lead to the production of winged offspring 66 for example densovirus infection has a negative impact on dysaphis plantaginea rosy apple aphid reproduction but contributes to the development of aphids with wings which can transmit the virus more easily to new host plants 67 additionally symbiotic bacteria that live inside of the aphids can also alter aphid reproductive strategies based on the exposure to environmental stressors 68 in the autumn host alternating heteroecious aphid species produce a special winged generation that flies to different host plants for the sexual part of the life cycle flightless female and male sexual forms are produced and lay eggs 69 some species such as aphis fabae black bean aphid metopolophium dirhodum rose grain aphid myzus persicae peach potato aphid and rhopalosiphum padi bird cherry oat aphid are serious pests they overwinter on primary hosts on trees or bushes in summer they migrate to their secondary host on a herbaceous plant often a crop then the gynoparae return to the tree in autumn another example is the soybean aphid aphis glycines as fall approaches the soybean plants begin to senesce from the bottom upwards the aphids are forced upwards and start to produce winged forms first females and later males which fly off to the primary host buckthorn here they mate and overwinter as eggs 52 ecology edit ant mutualism edit some species of ants farm aphids protecting them on the plants where they are feeding and consuming the honeydew the aphids release from the terminations of their alimentary canals this is a mutualistic relationship with these dairying ants milking the aphids by stroking them with their antennae c 70 although symbiotic the feeding behaviour of aphids is altered by ant attendance aphids attended by ants tend to increase the production of honeydew in smaller drops with a greater concentration of amino acids 71 macrosiphoniella yomogicola has green and red morphs in japan where lasius japonicus ants prefer the green morph which provides more honeydew and select for them their honeydew contains dopamine and makes the attendant ants act more aggressively against intruders the aphids are thought to be selfishly manipulating the ants to provide them better protection from predators and parasites 72 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 73 manage large herds of aphids that feed on roots of plants in the ant colony queens 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 70 some bees in coniferous forests collect aphid honeydew to make forest honey 29 an interesting variation in ant aphid relationships involves lycaenid butterflies and 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 since the caterpillars produce a pheromone which deceives the ants into treating them like ants and carrying the caterpillars into their nest once there the ants feed the caterpillars which in return produce honeydew for the ants when the caterpillars reach full size they crawl to the colony entrance and form cocoons after two weeks the adult butterflies emerge and take flight at this point the ants attack the butterflies but the butterflies have a sticky wool like substance on their wings that disables the ants jaws allowing the butterflies to fly away without being harmed 74 another ant mimicking gall aphid paracletus cimiciformis eriosomatinae has evolved a complex double strategy involving two morphs of the same clone and tetramorium ants aphids of the round morph cause the ants to farm them as with many other aphids the flat morph aphids are aggressive mimics with a wolf in sheep s clothing strategy they have hydrocarbons in their cuticle that mimic those of the ants and the ants carry them into the brood chamber of the ants nest and raise them like ant larvae once there the flat morph aphids behave like predators drinking the body fluids of ant larvae 75 an ant guards its aphids ants tending aphids ant extracting honeydew from an aphid ants tending aphids and collecting honeydew secreted a wrinkled solder beetle flies in and eats an aphid before being chased away by the ants bacterial endosymbiosis edit endosymbiosis with micro organisms is common in insects with more than 10 of insect species relying on intracellular bacteria for their development and survival 76 aphids harbour a vertically transmitted from parent to its offspring obligate symbiosis with buchnera aphidicola the primary symbiont inside specialized cells the bacteriocytes 77 five of the bacteria genes have been transferred to the aphid nucleus 78 the original association is estimated to have occurred in a common ancestor 280 to 160 million years ago and enabled aphids to exploit a new ecological niche feeding on phloem sap of vascular plants b aphidicola provides its host with essential amino acids which are present in low concentrations in plant sap 79 the metabolites from endosymbionts are also excreted in honeydew 80 the stable intracellular conditions as well as the bottleneck effect experienced during the transmission of a few bacteria from the mother to each nymph increase the probability of transmission of mutations and gene deletions 81 82 as a result the size of the b aphidicola genome is greatly reduced compared to its putative ancestor 83 despite the apparent loss of transcription factors in the reduced genome gene expression is highly regulated as shown by the ten fold variation in expression levels between different genes under normal conditions 84 buchnera aphidicola gene transcription although not well understood is thought to be regulated by a small number of global transcriptional regulators and or th...