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Plant damage and yield response to the Russian wheat aphid (Homoptera: Aphididae) on susceptible and resistant winter wheats in Colorado.

Plant damage and yield response to the Russian wheat aphid, Diuraphis noxia (Mordvilko), were evaluated on a susceptible (TAM 107) and a resistant (RWA E1) winter wheat, Triticum aestivum L., in three Colorado locations in the 1993 and 1994 crop years. Russian wheat aphid was more abundant on TAM 107 than on RWA E1. Russian wheat aphid days per tiller were greater at the higher infestation levels. Yield losses as a result of Russian wheat aphid infestation occurred most of the time with TAM 107 but rarely with RWA E1. Seed densities were reduced at higher infestation levels in TAM 107 at two locations. Russian wheat aphids per tiller had a negative relationship to yield in TAM 107 but not in RWA E1. In TAM 107 yield decreased as aphid densities increased, but yield remained constant regardless of initial aphid abundance on RWA E1 in all environments. Seed densities were reduced at higher infestation levels in TAM 107 at two locations. The resistance conferred by the Dn4 gene seems to be an effective management approach across a range of field conditions.

Animals↗

Comparison of bioassay techniques for determining baseline susceptibilities to imidacloprid for green apple aphid (Homoptera: Aphididae).

The susceptibility of a clone of green apple aphid, Aphis pomi (De Geer), to the neonicotinyl insecticide imidacloprid was determined by direct and indirect bioassay techniques. Aphid numbers were assessed on potted apple seedlings treated with various concentrations of imidacloprid, adults were dipped in test solutions as per the Food and Agriculture Organization protocol, or nymphs and adults were reared on treated apple leaf disks. Effective concentrations required to kill half of the test population (EC50) varied depending on the bioassay technique, ranging from as low as 0.064 ppm for first instars reared for 3 d on treated leaf disks to 1.79 ppm for adult apterae dipped in solutions of imidacloprid and held for 24 h on clean leaf disks. When imidacloprid was directly applied to aphids, mortality continued to increase over 3 d, but the difference was not statistically significant between day 2 (1.36 ppm) and day 3 (1.19 ppm). Toxicity of neonicotinyls to aphids is expressed rather slowly and primarily after oral ingestion. The effect of imidacloprid on reproduction of green apple aphid was also assessed for adult apterae reared on treated leaf disks. Contrary to previous reports, our results demonstrated that imidacloprid does not have a direct negative effect on the reproductive physiology of this species. Negative effects can mostly be attributed to the antifeedant activity of this compound and the protracted time to death. The results of this study contribute to a better understanding of the most suitable techniques for assessing aphid mortality after exposure to these new insecticides and provides a baseline susceptibility to imidacloprid for green apple aphid.

Animals↗

Field evaluation of emmer wheat-derived synthetic hexaploid wheat for resistance to Russian wheat aphid (Homoptera: Aphididae).

Broadening the genetic base for resistance to Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: Aphididae), in bread wheat, Triticum aestivum L., is desirable. To date, identified Russian wheat aphid resistance genes are either located to the D chromosomes or to rye translocation of wheat, and resistance derived from the A or B genomes of tetraploid Triticum spp. would therefore be highly beneficial. Fifty-eight synthetic hexaploid wheat, derived from interspecific crosses of Triticum dicoccum Schrank. and Aegilops tauschii (Coss.) Schmal. and their parents were evaluated for resistance to Russian wheat aphid under field conditions. Plots infested with aphids were compared with plots protected with insecticides. The T. dicoccum parents were highly resistant to Russian wheat aphids, whereas the Ae. tauschii parents were susceptible. Resistance levels observed in the synthetic hexaploids were slightly below the levels of their T. dicoccum parents when a visual damage scale was used. but no major resistance suppression was observed among the synthetics. Russian wheat aphid infestation on average reduced plant height and kernel weight at harvest in the synthetic hexaploids and the T. dicoccum parents by 3-4%, whereas the susceptible control 'Seri M82' suffered losses of above 20%. Because resistance in the synthetic hexaploid wheat is derived from their T. dicoccum parent, resistance gene(s) must be located on the A and/or B genomes. They must therefore be different from previously identified Russian wheat aphid resistance genes, which have all been located on the D genome of wheat or on translocated segments.

Animals↗

Aphid-induced defense responses in Mi-1-mediated compatible and incompatible tomato interactions.

The tomato Mi-1 gene confers resistance to three species of root-knot nematode and potato aphid. We studied changes in expression of jasmonic acid (JA)- and salicylic acid (SA)-dependent defense genes in response to potato and green peach aphids. We determined changes in three PR proteins, lipoxygenase and proteinase inhibitors I and II transcripts, locally and systemically in both compatible and incompatible interactions in tomato. Transcripts for PR-1 were detected earlier and accumulated to higher levels in the incompatible than in the compatible potato aphid/tomato interactions. The transcript profiles of the other genes were similar in compatible compared with incompatible interactions. Pin1 and Pin2 RNAs were detected early and transiently in both compatible and incompatible interactions. In tomato plants containing Mi-1, systemic expression of PR-1 and GluB was detected in both compatible and incompatible interactions at 48 h after infestations with either aphid. These results suggest that aphid feeding involves both SA and JA/ethylene plant defense signaling pathways and that Mi-1-mediated resistance might involve a SA-dependent signaling pathway. Potato aphid feeding generated reactive oxygen species in both compatible and incompatible interactions. However, a hypersensitive response was absent in the Mi-1-mediated resistance response to potato aphids. Reciprocal grafting experiments revealed that resistance is cell autonomous, and local expression of Mi-1 is required for Mi-1-mediated resistance against the potato aphid.

Animals↗

Changing partners in an obligate symbiosis: a facultative endosymbiont can compensate for loss of the essential endosymbiont Buchnera in an aphid.

Almost all aphids harbour an endosymbiotic bacterium, Buchnera aphidicola, in bacteriocytes. Buchnera synthesizes essential nutrients and supports growth and reproduction of the host. Over the long history of endosymbiosis, many essential genes have been lost from the Buchnera genome, resulting in drastic genome reduction and the inability to live outside the host cells. In turn, when deprived of Buchnera, the host aphid suffers retarded growth and sterility. Buchnera and the host aphid are often referred to as highly integrated almost inseparable mutualistic partners. However, we discovered that, even after complete elimination of Buchnera, infection with a facultative endosymbiotic gamma-proteobacterium called pea aphid secondary symbiont (PASS) enabled survival and reproduction of the pea aphid. In the Buchnera-free aphid, PASS infected the cytoplasms of bacteriocytes that normally harbour Buchnera, establishing a novel endosymbiotic system. These results indicate that PASS can compensate for the essential role of Buchnera by physiologically and cytologically taking over the symbiotic niche. By contrast, PASS negatively affected the growth and reproduction of normal host aphids by suppressing the essential symbiont Buchnera. These findings illuminate complex symbiont-symbiont and host-symbiont interactions in an endosymbiotic system, and suggest a possible evolutionary route to novel obligate endosymbiosis by way of facultative endosymbiotic associations.

Animals↗

Signal for potyvirus-dependent aphid transmission of potato aucuba mosaic virus and the effect of its transfer to potato virus X.

A British isolate of potato aucuba mosaic potexvirus (PAMV) was transmitted by aphids (Myzus persicae) which had fed previously on a source of potato Y potyvirus (PVY). Nucleotide sequence analysis of the PAMV coat protein gene indicated that amino acid residues 14 to 16 from the N terminus of the coat protein have the sequence DAG, which is also found in the coat proteins of potyviruses and is required for their aphid transmissibility. A recombinant virus isolate (TXPA7) was produced in which a segment of the coat protein gene of PAMV encoding the 40 N-terminal amino acids was inserted in the genome of potato X potexvirus (PVX) in place of the segment encoding the 28 N-terminal amino acids of PVX coat protein. This isolate, and a second similar recombinant (TXPA5) in which the DAG motif was changed to YTS, were mechanically transmissible to intact plants, in which they caused slightly milder symptoms than PVX. Particles of TXPA7 reacted in immunosorbent electron microscopy with PVX- and PAMV-specific antibodies and so were antigenically distinguishable from PAMV and PVX particles, which reacted only with their homologous antibody, and from TXPA5 particles, which reacted only with the PVX antibody. Recombinant TXPA7 was transmitted by aphids that had already fed on a source of PVY whereas TXPA5 and PVX were not. TXPA7 was not transmitted by aphids that had not fed on a PVY source. It is concluded that (i) the potyvirus-dependent aphid transmissibility of PAMV results from possession of a domain which includes the DAG motif and is located near the N terminus of the virus coat protein, and (ii) potyvirus-dependent aphid transmissibility can be conferred on PVX, a non-aphid-borne potexvirus, by substituting this domain for the N-terminal part of its coat protein.

Amino Acid Sequence↗

Posterior midgut and hindgut are both sites of acquisition of Cucurbit aphid-borne yellows virus in Myzus persicae and Aphis gossypii.

Members of the family Luteoviridae ('luteovirids') rely strictly on aphid vectors for plant-to-plant transmission. This interaction operates according to a persistent and circulative manner, which implies that the virions are being endocytosed and exocytosed across two epithelial barriers (alimentary tract and accessory salivary glands) in the vector's body. In several luteovirid-aphid vector species combinations, the route of virions in the insect has been investigated ultrastructurally by transmission electron microscopy (TEM). Here, we used TEM to follow the route of Cucurbit aphid-borne yellows virus (CABYV; genus Polerovirus) in its two efficient vector species, Myzus persicae and Aphis gossypii. We demonstrated that CABYV particles are acquired from the gut lumen to the haemocoel through two different sites in both aphid species, i.e. the posterior midgut (as for Beet western yellows virus in M. persicae) and the hindgut (as for Barley yellow dwarf virus complex in cereal aphids). This 'dual' tissue specificity of CABYV represents an original situation among viruses in the family Luteoviridae examined so far by TEM. A variety of virion-containing structures (e.g. clathrin-coated and tubular vesicles, endosome-like bodies) are found in intestinal cells of both types in both aphids. Release of virus particles from midgut and hindgut cells into the haemolymph was confirmed by immunotrapping using CABYV-specific antibodies. In accessory salivary glands, transport of CABYV virions across the cells was similar in each aphid species, and occurred by a transcytosis mechanism involving formation of tubular and coated vesicles before release of free virions in the salivary canal.

Animals↗

Aphid resistance in Medicago truncatula involves antixenosis and phloem-specific, inducible antibiosis, and maps to a single locus flanked by NBS-LRR resistance gene analogs.

Aphids and related insects feed from a single cell type in plants: the phloem sieve element. Genetic resistance to Acyrthosiphon kondoi Shinji (bluegreen aphid or blue alfalfa aphid) has been identified in Medicago truncatula Gaert. (barrel medic) and backcrossed into susceptible cultivars. The status of M. truncatula as a model legume allows an in-depth study of defense against this aphid at physiological, biochemical, and molecular levels. In this study, two closely related resistant and susceptible genotypes were used to characterize the aphid-resistance phenotype. Resistance conditions antixenosis since migratory aphids were deterred from settling on resistant plants within 6 h of release, preferring to settle on susceptible plants. Analysis of feeding behavior revealed the trait affects A. kondoi at the level of the phloem sieve element. Aphid reproduction on excised shoots demonstrated that resistance requires an intact plant. Antibiosis against A. kondoi is enhanced by prior infestation, indicating induction of this phloem-specific defense. Resistance segregates as a single dominant gene, AKR (Acyrthosiphon kondoi resistance), in two mapping populations, which have been used to map the locus to a region flanked by resistance gene analogs predicted to encode the CC-NBS-LRR subfamily of resistance proteins. This work provides the basis for future molecular analysis of defense against phloem parasitism in a plant model system.

Animals↗

An association between phlebotomine sandflies and aphids in the Peruvian Andes.

As the composition of natural sugars in the diet of adult sandflies (Diptera: Psychodidae: Phlebotominae) may affect the development of Leishmania (Kinetoplastida: Trypanosomatidae) in sandfly guts, and so play an important role in the epidemiology of leishmaniasis, there is increasing interest in the sources of sugars for wild sandflies. Advanced chromatography techniques have provided indirect evidence that wild sandflies feed on honeydew, a substance released by aphids (Hemiptera: Aphididae) when feeding. Our objective was to determine whether sandfly density can be influenced directly by the local density of aphids. Aphid density was determined by counting absolute numbers of aphids on alfalfa stems in Purisima Valley, Peru, where sandflies transmit Leishmania peruviana causing Andean cutaneous leishmaniasis (uta). Sandfly relative abundance was measured using sticky trap sampling repeatedly in alfalfa fields. Lutzomyia verrucarum accounted for 92% of the total sandflies collected. As there was a female bias in sandflies collected close to houses, only the numbers of male sandflies were used in analysis. Most of the adult aphids found feeding on alfalfa were either Therioaphis trifolii forma maculata (97%) or Acyrthosiphon pisum (3%). By regression analysis, a significant relationship was found between the density of Lu.verrucarum males and the density of adults of both aphid species. This is the first ecological study to support the hypothesis that aphid honeydew may be a source of sugar for sandflies.

Animals↗

The longest 18S ribosomal RNA ever known. Nucleotide sequence and presumed secondary structure of the 18S rRNA of the pea aphid, Acyrthosiphon pisum.

An EMBL4 recombinant phage which encodes one of the full length of the aphid ribosomal DNA has been isolated from the aphid genomic library. Determination of the complete nucleotide sequence of the aphid 18S rRNA gene revealed that it is 2469 bp with a G + C content of 59%. The aphid 18S rRNA gene studied here is the longest and has the highest G + C content among the 18S rRNA genes examined so far. Evidence provided by the S1 nuclease assay suggests that the aphid 18S rRNA gene examined in this study is not a pseudogene containing an insertion sequence. Based on the nucleotide sequence of the 18S rRNA gene, we constructed a presumed secondary-structure model of the aphid 18S rRNA. In the aphid 18S rRNA, the eucaryote-specific E21 and 41 region are supposed to be longer and more complex than the counterparts of other 18S rRNA.

Animals↗

The phytopathogen Dickeya dadantii (Erwinia chrysanthemi 3937) is a pathogen of the pea aphid.

Dickeya dadantii (Erwinia chrysanthemi) is a phytopathogenic bacterium causing soft rot diseases on many crops. The sequencing of its genome identified four genes encoding homologues of the Cyt family of insecticidal toxins from Bacillus thuringiensis, which are not present in the close relative Pectobacterium carotovorum subsp. atrosepticum. The pathogenicity of D. dadantii was tested on the pea aphid Acyrthosiphon pisum, and the bacterium was shown to be highly virulent for this insect, either by septic injury or by oral infection. The lethal inoculum dose was calculated to be as low as 10 ingested bacterial cells. A D. dadantii mutant with the four cytotoxin genes deleted showed a reduced per os virulence for A. pisum, highlighting the potential role of at least one of these genes in pathogenicity. Since only one bacterial pathogen of aphids has been previously described (Erwinia aphidicola), other species from the same bacterial group were tested. The pathogenic trait for aphids was shown to be widespread, albeit variable, within the phytopathogens, with no link to phylogenetic positioning in the Enterobacteriaceae. Previously characterized gut symbionts from thrips (Erwinia/Pantoea group) were also highly pathogenic to the aphid, whereas the potent entomopathogen Photorhabdus luminescens was not. D. dadantii is not a generalist insect pathogen, since it has low pathogenicity for three other insect species (Drosophila melanogaster, Sitophilus oryzae, and Spodoptera littoralis). D. dadantii was one of the most virulent aphid pathogens in our screening, and it was active on most aphid instars, except for the first one, probably due to anatomical filtering. The observed difference in virulence toward apterous and winged aphids may have an ecological impact, and this deserves specific attention in future research.

Animals↗

Genetics, physiology, and evolutionary relationships of the genus Buchnera: intracellular symbionts of aphids.

Evolutionary studies suggest that 200-250 million years ago an aphid ancestor was infected with a free-living eubacterium. The latter became established within aphid cells. Host and endosymbiont (genus Buchnera) became interdependent and unable to survive without each other. The growth of Buchnera became integrated with that of the aphids, which acquired the endosymbionts from their mothers before birth. Speciation of host lineages was paralleled by divergence of associated endosymbiont lineages, resulting in parallel evolution of Buchnera and aphids. Present day Buchnera retains many of the properties of its free-living ancestor, containing genes for proteins involved in DNA replication, transcription, and translation, as well as chaperonins and proteins involved in secretion, energy-yielding metabolism, and amino acid biosynthesis. Some of these processes are also observed in isolated endosymbiont cells. Genetic and physiological studies indicate that Buchnera can synthesize methionine, cysteine, and tryptophan and supply these amino acids to the aphid host. In the case of some fast-growing species of aphids, the overproduction of tryptophan by Buchnera involves plasmid-amplification of the gene coding for anthranilate synthase, the first enzyme of the tryptophan biosynthetic pathway. These recent studies provide a beginning in our understanding of Buchnera and its role in the endosymbiosis with aphids.

Animals↗

Low temperature acclimated populations of the grain aphid Sitobion avenae retain ability to rapidly cold harden with enhanced fitness.

In contrast to previous studies of rapid cold-hardening (RCH), which have investigated the responses of insects maintained under 'summer conditions' (20 degrees to 25 degrees C), this study focuses on the ability of low-temperature acclimated insects to undergo RCH. When the grain aphid Sitobion avenae Fabricus was low-temperature acclimated by rearing for three generations at 10 degrees C, the discriminating temperatures (temperature that results in approximately 20% survival after direct transfer from the rearing temperature to a sub-zero temperature for a period of 3 h), of first instar nymphs and adult aphids were -11.5 degrees and -12 degrees C, respectively. Maximum rapid cold-hardening was induced by cooling aphids at 0 degrees C for 2 h (nymphs) or 30 min (adults), resulting in survival at the respective discriminating temperatures increasing from 26% to 96% (nymphs) and 22% to 70% (adults). Cooling from 10 degrees to 0 degrees C at 1 degree, 0.1 degrees and 0.05 degrees C min-1 significantly increased survival of nymphs at the discriminating temperature, but not of adults. There were no ;ecological costs' associated with rapid cold-hardening at 0 degrees C, or with exposure of rapidly cold-hardened aphids to the discriminating temperatures; fecundity and longevity, in both nymphs and adults were either similar to control aphids or significantly increased. The study demonstrates that rapid cold-hardening ability is retained in aphids that have already undergone cold-acclimation, as would be the case in overwintering aphids. Both rapid cold-hardening and subsequent exposure at previously lethal temperatures can enhance fitness in surviving individuals.

Acclimatization↗

The impact of host plant on the abundance and function of symbiotic bacteria in an aphid.

The black-bean aphid Aphis fabae bears populations of coccoid symbiotic bacteria Buchnera spp. at 2.0-3.2 x 10(7)cells mg(-1)aphid mass and rod-shaped secondary symbionts of uncertain taxonomic affiliation at 0.1-0.6 x 10(7)cells mg(-1)aphid mass. Buchnera provides essential amino acids, supplementing the poor supply in the aphid diet of plant phloem sap. Comparison of the performance of A. fabae containing and experimentally deprived of their bacteria showed that the bacteria caused increased larval mass of aphids reared on Chenopodium album and Papaver dubium plants, but not when reared on Lamium purpureum. In the aphids reared on L. purpureum, the density of the bacteria, especially the secondary symbionts, was significantly elevated, and bacterial-mediated production of the essential amino acid threonine was reduced, even though the essential amino acid content of phloem exudates from L. purpureum had a low threonine content. It is proposed that the shortfall in threonine, possibly compounded by the high density of secondary symbionts, may contribute to the poor performance of the aphids on L. purpureum. This study offers the first evidence to suggest plant-mediated interference with the nutritional function of symbiotic bacteria in any phytophagous insect.

Amino Acids↗

Demography of soybean aphid (Homoptera: Aphididae) at summer temperatures.

Soybean aphid, Aphis glycines Matsumura, is now widely established in soybean, Glycine max L., production areas of the northern United States and southern Canada and is becoming an important economic pest. Temperature effect on soybean aphid fecundity and survivorship is not well understood. We determined the optimal temperature for soybean aphid growth and reproduction on soybean under controlled conditions. We constructed life tables for soybean aphid at 20, 25, 30, and 35 degrees C with a photoperiod of 16:8 (L:D) h. Population growth rates were greatest at 25 degrees C. As temperature increased, net fecundity, gross fecundity, generation time, and life expectancy decreased. The prereproductive period did not differ between 20 and 30 degrees C; however, at 30 degrees C aphids required more degree-days (base 8.6 degrees C) to develop. Nymphs exposed to 35 degrees C did not complete development, and all individuals died within 11 d. Reproductive periods were significantly different at all temperatures, with aphids reproducing longer and producing more progeny at 20 and 25 degrees C than at 30 or 35 degrees C. Using a modification of the nonlinear Logan model, we estimated upper and optimal developmental thresholds to be 34.9 and 27.8 degrees C, respectively. At 25 degrees C, aphid populations doubled in 1.5 d; at 20 and 30 degrees C, populations doubled in 1.9 d.

Animals↗

Determining the optimal timing of foliar insecticide applications for control of soybean aphid (Hemiptera: Aphididae) on soybean.

Field experiments were performed over 3 yr to examine the impact of insecticide application timing to control soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), populations and to prevent soybean yield losses. Experiments were conducted in early and late-planted soybean, Glycine max (L.) Merr. Insecticide applications were made based on soybean growth stages. In 2001, applications were made at V1, V3, R2, and R3 growth stages; in 2002 and 2003, applications were made at R2, R3, and R4 stages. Additional treatments consisted of an unsprayed control and a multiple spray treatment that received insecticide applications at 7-10-d intervals. Soybean aphid densities were recorded throughout the growing season, and yields were measured. Soybean aphid populations varied considerably across years and planting dates. In general, late-planted soybean exhibited higher aphid pressure than early planted soybean, and experiments in 2002 had lower aphid numbers than those in 2001 and 2003. The multiple spray treatment significantly increased yield over the control in four of the six experiments, the exceptions being 2002 late planted and 2003 early planted. This suggests that soybean aphid populations were not large enough to cause yield losses in these two experiments. The R3 spray treatment increased yield in three of the six experiments (2001 late planting, 2002 early planting, and 2003 late planting), the R2 spray treatment increased yield in two of six experiments (2001 and 2003 late plantings), and the V1 application increased yield over the control in the 2001 late-planted experiment. Results suggest that when aphid populations are high insecticide applications made at R2 and R3 plant stages are most effective in preventing yield loss.

Animals↗

Soldiers effectively defend aphid colonies against predators in the field

Morphologically specialized soldiers occur in more than 50 aphid species in the families Pemphigidae and Hormaphididae. To study the effectiveness of soldiers of the gall-forming aphid, Pemphigus spyrothecae Pass., in protecting their galls against natural levels of predation, we manipulated the proportions of soldiers and non-soldiers in sets of galls still attached to poplar trees in the field. Galls with 50 soldiers and 50 non-soldiers were approximately 10 times less likely to be attacked by predators than galls that contained 100 non-soldier aphids. There were significantly fewer live aphids, and significantly more dead aphids, in galls without soldiers than in galls protected either by soldiers or by being within a bag. There were no significant differences in the survival of aphids in galls protected by soldiers compared with those protected by bagging. The soldiers did not protect the galls against invasion by the cohabiting aphid Chaitophorus leucomelas Koch. These observations provide the first demonstration that soldiers are effective in defence against natural levels of predation under field conditions. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

A cost of alarm pheromone production in cotton aphids, Aphis gossypii.

The sesquiterpene, (E)-beta-farnesene, is used by many aphid species as an alarm pheromone to warn related individuals of predation. Disturbed cotton aphids, Aphis gossypii Glover, released (E)-beta-farnesene into the air as detected by solid phase microextraction and gas chromatography mass spectrometry (GC-MS). Solvent extracts of cotton aphids of various life stages and weights also were analyzed by GC-MS for sums of ions 69 and 93, which discriminated (E)-beta-farnesene from coeluting compounds. Aphids of all life stages and sizes reared on cotton plants in both an environmental chamber and glasshouse contained (E)-beta-farnesene in amounts ranging from 0.1 to 1.5 ng per individual. The quantities of (E)-beta-farnesene in aphids increased in relation to increasing body weight, and variation in individual weights explained about 82% of the variation in alarm pheromone. However, the concentrations (ng/mg fresh weight) declined exponentially with increasing body weight. These findings indicate that aphid nymphs try to compensate for their smaller size by producing relatively more pheromone per weight than adults but still cannot approach an evolutionary optimal load, as assumed in adults with the greatest total amounts. This suggests that young aphids need to balance costs of growth and maturation with costs of producing the alarm pheromone.

Animals↗