Arianne J. Cease

Heavy Livestock Grazing Promotes Locust Outbreaks by Lowering Plant Nitrogen Content

Locust Heaven Locust outbreaks have severe consequences for agriculture, but the conditions that promote an outbreak are unknown. Cease et al. (p. 467) investigated aspects of the locust diet and found that increased nitrogen content of cereal grasses reduced the size and viability of a herbivorous locust species. This locust prefers low N plants, which result from heavy grazing by livestock and erosion. High-protein plants inhibit locust swarming, which explains why grazed systems are more prone to outbreaks. Current paradigms generally assume that increased plant nitrogen (N) should enhance herbivore performance by relieving protein limitation, increasing herbivorous insect populations. We show, in contrast to this scenario, that host plant N enrichment and high-protein artificial diets decreased the size and viability of Oedaleus asiaticus, a dominant locust of north Asian grasslands. This locust preferred plants with low N content and artificial diets with low protein and high carbohydrate content. Plant N content was lowest and locust abundance highest in heavily livestock-grazed fields where soils were N-depleted, likely due to enhanced erosion. These results suggest that heavy livestock grazing and consequent steppe degradation in the Eurasian grassland promote outbreaks of this locust by reducing plant protein content.

How Locusts Breathe

Insect tracheal-respiratory systems achieve high fluxes and great dynamic range with low energy requirements and could be important models for bioengineers interested in developing microfluidic systems. Recent advances suggest that insect cardiorespiratory systems have functional valves that permit compartmentalization with segment-specific pressures and flows and that system anatomy allows regional flows. Convection dominates over diffusion as a transport mechanism in the major tracheae, but Reynolds numbers suggest viscous effects remain important.

Are color or high rearing density related to migratory polyphenism in the band-winged grasshopper, Oedaleus asiaticus?

Locusts represent an impressive example of migratory polyphenism, with high densities triggering a switch from a solitarious, shorter dispersal range, and sometimes greenish phenotype to a gregarious and sometimes darker form exhibiting behavioral, morphological and physiological traits associated with long-distance migratory swarms. While such polyphenism has been well documented in Locusta migratoria and Schistocerca gregaria, the extent to which other grasshoppers exhibit this type of migratory polyphenism is unclear. Anecdotally, the Chinese grasshopper, Oedaleus asiaticus, forms migratory swarms comprised mostly of a darker, brown-colored morph, but also exhibits a non-migratory green-colored morph that predominates at low densities. In a population in Inner Mongolia not currently exhibiting migratory swarms, we found that while green and brown O. asiaticus are found concurrently across our sampled range, only brown grasshoppers were found in high densities. Differences between field-collected brown and green forms matched some but not key predictions associated with the hypothesis that the brown form is morphologically and physiologically specialized for gregarious migration. Controlling for body mass, brown forms had more massive thoraxes, abdomens and legs, and higher metabolic rates, but not more flight muscle or lipid stores. Further, the brown and green grasshoppers did not differ in gregarious behavior, and neither would fly in multiple lab and field trials. Lab or field-rearing at high densities for one-to-multiple juvenile instars caused grasshoppers to exhibit some morphological traits predicted to benefit migration (larger wings and a shift in relative mass from abdomen to thorax), but did not change color or induce flight behavior. One hypothesis to explain these data is that a migratory form of O. asiaticus is partially triggered by high field densities, but that existing ecological conditions blocked full expression of such traits (and outbreak swarms). Alternatively, color variation in this species may more tightly linked to other functions in this species such as crypsis or disease resistance, and mechanisms other than late-juvenile rearing density (e.g. genetic variation, maternal effects) may be more critical for promoting variation in color and/or migratory polyphenism.

Caterpillars selected for large body size and short development time are more susceptible to oxygen-related stress

Recent studies suggest that higher growth rates may be associated with reduced capacities for stress tolerance and increased accumulated damage due to reactive oxygen species. We tested the response of Manduca sexta (Sphingidae) lines selected for large or small body size and short development time to hypoxia (10 kPa) and hyperoxia (25, 33, and 40 kPa); both hypoxia and hyperoxia reduce reproduction and oxygen levels over 33 kPa have been shown to increase oxidative damage in insects. Under normoxic (21 kPa) conditions, individuals from the large-selected (big-fast) line were larger and had faster growth rates, slightly longer developmental times, and reduced survival rates compared to individuals from a line selected for small size (small-fast) or an unselected control line. Individuals from the big-fast line exhibited greater negative responses to hyperoxia with greater reductions in juvenile and adult mass, growth rate, and survival than the other two lines. Hypoxia generally negatively affected survival and growth/size, but the lines responded similarly. These results are mostly consistent with the hypothesis that simultaneous acquisition of large body sizes and short development times leads to reduced capacities for coping with stressful conditions including oxidative damage. This result is of particular importance in that natural selection tends to decrease development time and increase body size.

Grasshoppers Regulate N:P Stoichiometric Homeostasis by Changing Phosphorus Contents in Their Frass

Nitrogen (N) and phosphorus (P) are important limiting nutrients for plant production and consumer performance in a variety of ecosystems. As a result, the N:P stoichiometry of herbivores has received increased attention in ecology. However, the mechanisms by which herbivores maintain N:P stoichiometric homeostasis are poorly understood. Here, using a field manipulation experiment we show that the grasshopper Oedaleus asiaticus maintains strong N:P stoichiometric homeostasis regardless of whether grasshoppers were reared at low or high density. Grasshoppers maintained homeostasis by increasing P excretion when eating plants with higher P contents. However, while grasshoppers also maintained constant body N contents, we found no changes in N excretion in response to changing plant N content over the range measured. These results suggest that O. asiaticus maintains P homeostasis primarily by changing P absorption and excretion rates, but that other mechanisms may be more important for regulating N homeostasis. Our findings improve our understanding of consumer-driven P recycling and may help in understanding the factors affecting plant-herbivore interactions and ecosystem processes in grasslands.

Dietary phosphate affects food selection, post-ingestive phosphorus fate, and performance of a polyphagous herbivore

ABSTRACT Comparisons of the carbon, nitrogen and phosphorus (P) content of plants and insect herbivores suggests that P limitation and herbivore foraging to balance P intake could be common. However, the lack of synthetic diets for testing the effects of lower ranges of dietary P has been a major impediment to experimental assessment of the ecological importance of, and physiological responses to, P limitation for terrestrial herbivores. We manipulated dietary P content (%P) over its observed range in terrestrial foliage using artificial diets containing near-optimal content of other nutrients for the grasshopper Schistocerca americana. Over much of the ecologically relevant range, when consuming single diets over a lifetime, higher P stimulated growth rates and increased survival, with an optimal dietary %P of 0.25–0.50% when measured throughout development. Excessive dietary P (1%) reduced growth and survival. However, with only short-term (3 day) confinement to single diets, dietary P had no effect on food consumption or growth rates. During these short exposures, fifth (but not third) instar hoppers increased the proportion of P excreted relative to P assimilated as dietary P increased. Target experiments demonstrated that, when given a choice, grasshoppers select among foods to attain a P intake target of 0.6%. These data suggest that P limitation could be common for terrestrial insect herbivores and that they can exhibit ingestive and post-ingestive mechanisms to attain sufficient but not excessive P. Summary: Phosphorus limitation may be common for terrestrial herbivores; in response, grasshoppers can modulate P assimilation over a moderate range but extreme and/or long-term consumption of low-P or indeed even high-P food decreases growth and survival.

Nutritional imbalance suppresses migratory phenotypes of the Mongolian locust (Oedaleus asiaticus)

For many species, migration evolves to allow organisms to access better resources. However, the proximate factors that trigger these developmental changes, and how and why these vary across species, remain poorly understood. One prominent hypothesis is that poor-quality food promotes development of migratory phenotypes and this has been clearly shown for some polyphenic insects. In other animals, particularly long-distance bird migrants, it is clear that high-quality food is required to prepare animals for a successful migration. We tested the effect of diet quality on the flight behaviour and morphology of the Mongolian locust, Oedaleus asiaticus. Locusts reared at high population density and fed low-N grass (performance-enhancing for this species) had enhanced migratory morphology relative to locusts fed high-N grass. Furthermore, locusts fed synthetic diets with an optimal 1 : 2 protein : carbohydrate ratio flew for longer times than locusts fed diets with lower or higher protein : carbohydrate ratios. In contrast to the hypothesis that performance-degrading food should enhance migration, our results support the more nuanced hypothesis that high-quality diets promote development of migratory characteristics when migration is physiologically challenging.

The impact of nitrogen enrichment on grassland ecosystem stability depends on nitrogen addition level

Increasing atmospheric nitrogen (N) deposition may affect plant biodiversity, subsequently altering ecosystem stability. While a few studies have explored how simulated N deposition affects community stability and its underlying mechanisms, the experimental levels of N addition used are usually higher than current and future N deposition rates. Thus, their results could produce highly uncertain predictions of ecosystem function, especially if the responses to N deposition are nonlinear. We conducted a manipulative experiment that simulated elevated atmospheric N deposition with several N addition levels to evaluate the effect of N deposition on ecosystem stability and its underlying mechanisms in a semiarid grassland in northern China. Here we show that N addition altered community diversity, reducing species richness, evenness, diversity and dominance. In addition, we found that N addition at current N deposition levels had no significant impact on community stability. In contrast, N addition at levels from 4.6 to 13.8gNm-2yr-1 significantly decreased community stability, although community stability for the 13.8gNm-2yr-1 treatment was higher than that for the 4.6gNm-2yr-1 treatment. These results indicate that the response of community stability to N enrichment is nonlinear. This nonlinear change in community stability was positively correlated with species asynchrony, species richness, and species diversity as well as the stability of dominant species and the stability of the grass functional group. Our data suggest a need to re-evaluate the mechanisms responsible for the effects of N deposition on natural ecosystem stability across multiple levels of N enrichment and that additional experimentation with gradients of N loads more similar to future atmospheric N deposition rates is needed.