The preference–performance hypothesis predicts that moth behaviour links plant variations with caterpillar attack and distribution, and the plant‐age hypothesis states that specialist herbivores are more successful in exploring younger plant tissue. We integrated these predictions to investigate underlying mechanisms by which moths and caterpillars of Plutella xylostella L. (Lepidoptera: Plutellidae) track and exploit within‐plant variability of leaf age and stratification. We measured leaf proteins, glucosinolates and fibre, as well as larval choice, developmental performance, and moth oviposition preference with regard to leaf age classes (young, mature and senescent) of three varieties (collard, cauliflower and cabbage) of the main host plant Brassica oleracea L. Larvae consistently fit the prediction that specialist herbivores prefer and perform better on young, upper leaves that have the highest protein level, despite the highest content of defence compounds. Conversely, moths laid more eggs on fibrous and less nutritious leaves from the lower and senescent stratum. We argue that the leaf stratification of host plants imposes conflicting selective pressures concerning offspring feeding and protection on adult females. If egg mortality is catastrophic on the upper nutritious leaves in a particular microclimatic context (e.g. sun, heat, winds, drought or rain‐washing), then oviposition preference will remain for the suboptimal lower and senescent leaves. The ability of larvae to spread upwards over the plant to access the more nutritious leaf stratum is critical when eggs are preferentially laid on the protective low‐quality leaves.
The present study aimed to verify (1) whether seasonal increases in neotropical populations of Plutella xylostella are directly provoked by regular influxes of migrants, and (2) whether temporal variation in food availability is the ecological process behind such predictable events. Over 3 years, plants that P. xylostella prefers were cultivated and irrigated in order to provide a continuous and abundant supply of food. Nevertheless, seasonal oscillations in the population of the herbivore still occurred. The hypothesis of seasonal availability of host plants could not explain the population pattern. In April, when the insect was practically extinct from the area, an artificial infestation (immigration) with 10,000 pupae established a precocious population. Therefore, the start of the natural cycles of population growth, during July–August, seems to be due to external factors, rather than an improvement in local conditions for resident individuals. In the beginning of the natural cycles, the increase in the density of adults significantly preceded the increase in immatures. Plutella xylostella does not diapause, and therefore immigration is the proximate cause of the seasonal population increases. Hypotheses about local factors are suggested to explain the decreasing phase of the predictable population cycles.
Soil amendment with Silicon (Si) can increase plant resistance against insect herbivores, but the underlying mechanisms remain unclear. The mechanical resistance hypothesis (MRH) states that Si accumulated in epidermal cells directly and passively protects against herbivores by creating a mechanical barrier. The physiological resistance hypothesis (PRH) states that Si enhances resistance by activating plant biochemical and physiological processes. We tested both hypotheses by manipulating Si fertilization of the Si non‐accumulator collard, Brassica oleracea L. cv. acephala (Brassicaceae). Then, we assessed functional and ultrastructural plant responses and the developmental and reproductive performance of the leaf‐chewing larvae of the diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), and the sap‐sucking cabbage aphid, Brevicoryne brassicae L. (Hemiptera: Aphididae). There was a 20% increase in leaf Si content. Silicon deposition in epidermal cells was identified by confocal microscopy and directly coincided with lower performance of P. xylostella, but did not affect B. brassicae. On the other hand, we found no unequivocal evidence that Si‐mediated changes in primary and secondary metabolism improved plant resistance against the insects. Negative mechanical effects of Si on the insects may have masked beneficial effects of increased water, nitrogen, and mineral contents in Si‐treated collards. Silicon did not change leaf contents of hemicellulose, cellulose, and lignin. Although Si‐mediated increases in leaf glucosinolates (GLS) correlated with lower larval performance and higher oviposition preference of P. xylostella, both P. xylostella and B. brassicae are highly specialized in overcoming such secondary metabolites. Thus, mechanical resistance may have impaired P. xylostella, rather than the Si‐mediated increase in GLS. We suggest that the PRH may depend on the degree of insect feeding specialization, so that toxic Si‐mediated defenses may be more efficient against unadapted polyphagous herbivores. For them, a toxic barrier may be added to the mechanical resistance.
An initial destabilization of functions triggered by drought stress in plants is followed by acclimatization and acquisition of tolerance; however, knowledge remains limited on drought-mediated changes in plant quality for herbivores. We tested whether a water-stressed fast-growing plant negatively affects host-specialist insects in both sap-sucking and leaf-chewing feeding guilds. Collards (Brassica oleracea var. acephala) were grown in wellwatered, slightly water-stressed and severely water-stressed conditions. Decreasing soil moisture adversely affected plant development, assessed as a reduction in leaf number and size, stomatal size and relative water content. Severely stressed plants had less fiber and glucosinolates; however, they showed more total nitrogen and lipids. Larval survival, pupal weight, reproductive rate (Ro) and rate of population growth (r) were lower when the leaf-chewing Plutella xylostella was reared with severely stressed collards. In multiple-choice tests, moths laid fewer eggs on leaf discs of collard that were exposed to drought. The fecundity of the sap-sucking Brevicoryne brassicae was higher and the development of alates was lower when insects were fed on plants kept in well-watered regime as compared to slight-stress and severe-stress. Despite higher nitrogen content and fewer glucosinolates, a higher level of leaf surface wax in severely stressed collards possibly decreased food quality for both herbivores. Thus, hostspecific herbivores of different guilds showed similar responses to drought-stressed, fast-growing plants. Waterstressed crops could discourage the attack of specialist insects, but the intensity of the stress that is required to achieve this effect will greatly reduce crop production, in terms of plant growth or foliage increment.
Performance de um Inseto Oligófago em Relação à Idade da Planta Hospedeira RESUMO -A performance de insetos folívoros monófagos e oligófagos deveria ser afetada negativamente pelo envelhecimento das suas plantas hospedeiras. Plutella xylostella L. (Lepidoptera: Yponomeutidae), uma especialista em Cruciferae, foi criada em laboratório com folhas destacadas de repolho (Brassica oleraceae var. capitata). Foram utilizadas a quarta folha expandida de plantas de diferentes idades mais as folhas dos três estratos da planta madura. O envelhecimento da planta aumentou a mortalidade preimaginal e reduziu a taxa de desenvolvimento das larvas, o peso das pupas e a fecundidade dos adultos. A taxa reprodutiva líquida (Ro) e a taxa intrínseca de crescimento populacional (r) diminuíram com o envelhecimento da planta. Nenhuma das categorias de folhas da planta madura mostrou-se qualitativamente superior às folhas de plantas mais jovens. Se em condições de campo existir uma correlação positiva entre preferencia de oviposição e performance da prole, espera-se uma tendência temporal decrescente na intensidade de ataque de larvas ao longo do desenvolvimento da planta. PALAVRAS-CHAVE: Desenvolvimento larval, sucesso reprodutivo, fenologia da planta, Plutella xylostella, LepidopteraABSTRACT -The performance of monophagous and oligophagous herbivore insects should be negatively influenced by the ageing of their host plants. Plutella xylostella L. (Lepidoptera: Yponomeutidae), a specialist in Cruciferae, was reared in the laboratory on detached cabbage leaves (Brassica oleraceae var. capitata). We used fourth expanded leaves from differently aged plants, and leaves from three strata of the mature plant. Plant ageing increased pre-imaginal mortality and reduced larval developing rate, pupae weight, and fecundity. The insect net reproductive rate (Ro) and intrinsic rate of population growth (r) decreased as plant aged. No leaf category of mature plants was qualitatively superior to leaves from younger plants. If there exist a positive correlation between oviposition preference and offspring performance, in field conditions, a decreasing temporal trend in the intensity of attack along the host plant development can be expected.
Plant defence mechanisms can reduce the digestive enzyme activity of insect pests. The aim of this study was to determine the relationship between the production of proteinase inhibitors, lipoxygenase and polyphenol oxidase activity in Coffea arabica (Catuai IAC 15) plants, and the digestive enzyme activity in the pest Leucoptera coffeella (Lepidoptera: Lyonetiidae) after feeding on the plant. The production of proteinase inhibitors was evaluated with L‐BApNA as a substrate. We studied lipoxygenase activity with linoleic acid and polyphenol oxidase activity with catechol substrates, in coffee plants damaged (T1) and not damaged (T2) by L. coffeella. L. coffeella digestive enzyme activity was verified by trypsin‐like (substrate l‐BApNA and l‐TAME), chymotrypsin‐like (BTpNA and ATEE), cysteine proteases (l‐BApNA) and total protease (azocasein). Proteinase inhibitor production and lipoxygenase and polyphenol oxidase activity in C. arabica increases (P ≤ 0.05) with L. coffeella damage. Our results provide important information that these enzymatic activities may play a role in plant defence processes in C. arabica. Trypsin‐like activity increases, whereas chymotrypsin‐like and cysteine protease activity decrease in the midgut of L. coffeella, which acts as a defence mechanism.
Purification of active trypsin in the digestive process of insects is essential for the development of potent protease inhibitors (PIs) as an emerging pest control technology and research into insect adaptations to dietary PIs. An important aspect is the presence of proteolytic microorganisms, which contribute to host nutrition. Here, we purified trypsins produced by bacteria Bacillus cereus, Enterococcus mundtii, Enterococcus gallinarum, and Staphylococcus xylosus isolated from the midgut of Anticarsia gemmatalis. The trypsins had a molecular mass of approximately 25 kDa. The enzymes showed increased activity at 40°C, and they were active at pH values 7.5-10. Aprotinin, bis-benzamidine, and soybean Kunitz inhibitor (SKTI) significantly inhibited trypsin activity. The l-1-tosyl-amido-2-phenylethylchloromethyl ketone (TPCK), pepstatin A, E-64, ethylenediamine tetraacetic acid, and calcium ions did not affect the enzyme activity at the concentrations tested. We infer the purified trypsins do not require calcium ions, by which they differ from the trypsins of other microorganisms and the soluble and insoluble trypsins characterized from A. gemmatalis. These data suggest the existence of different isoforms of trypsin in the velvetbean caterpillar midguts.
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