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.
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.
It was postulated that Si‐mediated plant resistance to herbivory changes with soil water status, increasing when plants are under drought stress. We subjected collard (Brassica oleracea) to such soil variables and assessed plant responses and effects on the leaf‐chewing larvae of Plutella xylostella and the sap‐sucking aphid Brevicoryne brassicae. Silicon accumulated in collard leaves independently of soil water conditions, but it influenced mainly drought‐stressed plants. Silicon suppressed harmful effects of drought on leaf and root length and raised leaf water content and stomatal size to the same conditions of well‐watered plants. Drought stress reduced hemicellulose and cellulose, but Si did not influence them or lignin. Combination of drought and Si increased total and soluble leaf nitrogen. Drought decreased total glucosinolates, but Si increased such defence metabolites to similar concentrations that were found in well‐watered plants. Nutritional changes mediated by drought and Si in fibre, leaf water content, soluble nitrogen and glucosinolates did not increase insect performance in any feeding guild. Instead, caterpillars performed worse in drought‐stressed or Si‐treated collards, mainly in plants under combined conditions. Silicon improved plant resistance to drought and herbivore stresses.
The local population dynamics of insect herbivores in ephemeral patches of short-lived plants are poorly known. We investigated whether a specialist and a generalist caterpillar exhibit contrasting temporal patterns of attack during plant development and also assessed bottom-up forces related to plant ontogeny that govern such population trends. Immature stages of the polyphagous Trichoplusia ni (Hübner) and the oligophagous Plutella xylostella (L.) were sampled throughout the development of cabbage (Brassica oleracea L. var. capitata L.) crops. We measured protein and glucosinolate contents and insect performance with regard to plant age and leaf strata. The populations of both caterpillar species changed in close parallel throughout plant development, and a nonlinear temporal pattern of egg laying was reproduced in sequential population patterns of the larval stages until pupation. Reduced protein availability and insect performance coincided with a decline in egg laying and subsequent larval abundance in mature plants. By standardizing the plant size, we found that young and nutritious plants support proportionately more insects than large and mature plants. In our models of the population oscillations, the interaction between plant size and quality provided a strong causal explanation for the densities of both oligophagous and polyphagous caterpillars. Patches of fast-growing herbaceous plants are very common worldwide in the form of crop fields, and a generalized temporal pattern of attack may be widespread among caterpillars, regardless of their feeding specialization. Our results highlight the role of bottom-up forces in shaping the population dynamics of caterpillars in such systems.
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