Protection via parasitism: Datura odors attract parasitoid flies, which inhibit Manduca larvae from feeding and growing but may not help plants.

Insect carnivores frequently use olfactory cues from plants to find prey or hosts. For plants, the benefits of attracting parasitoids have been controversial, partly because parasitoids often do not kill their host insect immediately. Furthermore, most research has focused on the effects of solitary parasitoids on growth and feeding of hosts, even though many parasitoids are gregarious (multiple siblings inhabit the same host). Here, we examine how a gregarious parasitoid, the tachinid fly Drino rhoeo, uses olfactory cues from the host plant Datura wrightii to find the sphingid herbivore Manduca sexta, and how parasitism affects growth and feeding of host larvae. In behavioral trials using a Y-olfactometer, female flies were attracted to olfactory cues emitted by attacked plants and by cues emitted from the frass produced by larval Manduca sexta. M. sexta caterpillars that were parasitized by D. rhoeo grew to lower maximum weights, grew more slowly, and ate less of their host plant. We also present an analytical model to predict how tri-trophic interactions change with varying herbivory levels, parasitization rates and plant sizes. This model predicted that smaller plants gain a relatively greater benefit compared to large plants in attracting D. rhoeo. By assessing the behavior, the effects of host performance, and the variation in ecological parameters of the system, we can better understand the complex interactions between herbivorous insects, the plants they live on and the third trophic level members that attack them.

Interactive effects of rearing temperature and oxygen on the development of Drosophila melanogaster.

Although higher temperatures strongly stimulate ectothermic metabolic rates, they only slightly increase oxygen diffusion rates and decrease oxygen solubility. Consequently, we predicted that insect gas exchange systems would have more difficulty meeting tissue oxygen demands at higher temperatures. In this study, Drosophila melanogaster were reared from egg to adult in hyperoxic (40%), hypoxic (10%), and normoxic (21%) conditions and in temperatures ranging from 15 degrees -31.5 degrees C to examine the interactive effect of temperature and oxygen on development. Hyperoxia generally increased mass and growth rate at higher rearing temperatures. At lower rearing temperatures, however, hyperoxia had a very small effect on mass, did not affect growth rate, and lengthened time to eclosion. Relative to normoxia, flies reared in hypoxic conditions were generally smaller (mass and thorax length), had longer eclosion times, slower growth rates, and reduced survival. At cooler temperatures, hypoxia had relatively modest or nonsignificant effects on development, while at higher temperatures, the effects of hypoxia were large. These results suggest that higher temperatures reduce oxygen delivery capacity relative to tissue oxygen needs, which may partially explain why ectotherms are smaller when development occurs at higher temperatures.

Contrasting responses to desiccation and starvation by eggs and neonates of two lepidoptera.

We examined the effects of desiccation on eggs and first-instar larvae of two species of Lepidoptera, Grammia geneura (Arctiidae) and Manduca sexta (Sphingidae). Grammia geneura occurs primarily in grasslands and savannas of the southwestern United States; M. sexta co-occurs with G. geneura but also is cosmopolitan across much of the Western Hemisphere. Eggs of G. geneura exposed to 0% relative humidity (RH) lost water much less rapidly (7.6 microg d(-1); 2.4% d(-1)) than did eggs of M. sexta (79.5 microg d(-1); 5.7% d(-1)). Eggs of both species survived at rates exceeding 75% at both 0% and 85% RH. Neonates of the two species responded differently to desiccation and starvation. In 85% RH, larval G. geneura survived at high rates (>80%) without access to food or water up to day 17, and in 0% RH, they survived at rates exceeding 50% through the first 10 d. Larvae at 0% RH lost mass very slowly (7.2 microg d(-1); 2.9% d(-1)), which was attributable both to low rates of water loss and to an ability to reduce metabolic rate to low levels. Larval M. sexta, in contrast, had rates of mortality that were much higher: after 1 d, fewer than 30% were alive in either group, and by about 1.5 d, all were dead. Neonate M. sexta also lost mass much more rapidly at 0% RH, about 329 microg d(-1). Water from metabolism appeared to contribute significantly to the water budget of G. geneura but not of M. sexta. These data show that G. geneura and M. sexta can inhabit similar macroclimates via remarkably different physiologies.

The beneficial acclimation hypothesis versus acclimation of specific traits: physiological change in water-stressed Manduca sexta caterpillars.

Do organisms make beneficial physiological adjustments in response to environmental change? We examined this question by measuring the effects of short-term (12-36 h) and long-term (larval lifetime) hydric stress on the tobacco hornworm, Manduca sexta. Larvae were reared from the first instar on low-water (69%) or high-water (80%) artificial diets and then transferred early in the fifth instar to the same or opposite diet (2x2 design). Within the subsequent 36 h, we measured 24-h growth rates and three primary determinants of the water budget: water gain via consumption and water loss via evaporation and defecation. Larvae preexposed to low-water diet grew less rapidly on low-water diet than those switched acutely to low-water diet from high-water diet, showing that larvae preexposed to a particular environment do not necessarily acclimate beneficially to that environment. Our data on water fluxes to and from larvae, however, strongly suggest that water-stressed larvae did make beneficial physiological adjustments. Larvae responded to short-term hydric stress by minimizing rates of water excretion, primarily by increasing rates of rectal water absorption. Larvae responded to chronic water stress by significantly reducing rates of evaporative water loss; they also showed additional reductions in fecal water excretion, but these decreases were due to lowered consumption and not to further increases in rate of rectal water absorption. This mismatch between maladaptive acclimation of organismal performance and beneficial adjustment of suborganismal traits can be reconciled by recognizing that organismal physiology is hierarchical: fitness-related performance traits represent the aggregate outcome of numerous, more mechanistic physiological traits. Although chronic exposure to an environment may depress the aggregate effect of these mechanistic traits on performance, organisms are not precluded from making beneficial adjustments to individual traits contributing to performance.

Patterns and mechanisms of growth of fifth-instar Manduca sexta caterpillars following exposure to low- or high-protein food during early instars.

For many insect herbivores, variation in protein availability is a pervasive part of the environment. I explore how variable protein availability affects growth rates of fifth-instar Manduca sexta caterpillars and how growth is related to behavior and physiology. Groups of larvae were reared on low- or high-protein artificial diets (5.9% and 17.7% casein by dry weight, respectively) and then transferred in the fifth instar to the same or opposite diet. During or after the 24-h period following transfer, I measured growth rate, consumption rate, growth efficiency, midgut proteolytic activity, and masses of midgut contents and tissues. Fifth-instar caterpillars reared in earlier instars on high-protein diet grew about 20% more rapidly over 24 h than did caterpillars reared on low-protein diet. This growth pattern appears to be caused by differences in consumption and growth efficiency: caterpillars reared on high protein consumed more food, and used it more efficiently, than did caterpillars reared on low-protein diet. Over the short term (24 h), in contrast, fifth instars that received low-protein diet grew as rapidly as caterpillars that received high-protein diet. Increased (compensatory) consumption appears to be the primary mechanism by which caterpillars consuming low-protein food maintained growth rates.

Effects of dietary protein concentration on L-proline transport by Manduca sexta midgut.

Foliar protein concentrations vary widely. Phytophagous insects may respond to this variation by regulating the rate of absorption of protein breakdown products. Using fifth-instar Manduca sexta reared on artificial diets, we examined whether the rate of L-proline transport across the posterior midgut was affected by dietary protein concentration. Caterpillars were reared to the fifth instar on a low- or high-protein diet and then transferred to the same or opposite diet for 24 h prior to the flux measurements. Posterior midguts were mounted in Ussing chambers, and both potential difference and (14)C-proline flux across the tissue were measured during two-hour experiments. Midguts transported proline from lumen to hemolymph at about 6.5 &mgr;mol/cm(2)/h and from hemolymph to lumen at less than 0.43 &mgr;mol/cm(2)/h. Only a small fraction was oxidized (0.58 &mgr;mol/cm(2)/h), and proline was not converted to another compound while crossing the midgut epithelium. Neither proline transport rates nor potential differences across the epithelium differed significantly among treatment groups, suggesting that larval M. sexta do not regulate proline transport in response to dietary protein variation.

Thermal sensitivity of growth and feeding in Manduca sexta caterpillars.

We explore how the thermal sensitivity of organismic performance emerges from the thermal sensitivity of the underlying component processes involved, using growth and feeding of Manduca sexta caterpillars as a model system. We measured thermal performance curves for the short-term rates of growth, consumption, protein (casein) digestion, amino acid (methionine) uptake, and respiration in fifth-instar caterpillars over a biologically realistic temperature range from 14 degrees to 42 degrees C. Growth and consumption rates increased between 14 degrees and 26 degrees C, reached a maximum value near 34 degrees C, and declined rapidly above 38 degrees C. In contrast, protein digestion rate and respiration rate increased monotonically over the entire temperature range, and amino acid uptake rate increased with temperatures up to 38 degrees C and then leveled off between 38 degrees and 42 degrees C. These results suggest that the shape and position of the thermal performance curve for growth rate--in particular the maximum at 34 degrees C and rapid decline above 38 degrees C--was most closely correlated with the thermal sensitivity of consumption rate; the declining growth performance above 38 degrees C was not associated with declines in digestion or uptake rates or with accelerated respiration rates at these temperatures.

Egg-Mass Gel of Melanochlamys diomedea (Bergh) Protects Embryos From Low Salinity.

Many opisthobranch gastropods embed their embryos in gelatinous egg masses; however, the functions of gel are not well known. We analyze the hypothesis that egg-mass gel protects embryos from salinity change. Using egg masses of Melanochlamys diomedea, we found that experimental removal of gel decreased the ability of embryos to survive osmotic stress. We evaluate several possible protective mechanisms by estimating osmotic influx of water into egg masses and by modeling salt efflux from an egg mass. On immersion in low-salinity water, egg masses lost roughly 23% of their mass, indicating that osmotic influx of water did not occur. Therefore, the principal route of salinity change within the egg mass is probably salt efflux. The model suggests that this efflux occurs quite slowly even when ambient salinity changes rapidly. Slow salinity change may be less stressful for embryos because the mechanisms that regulate cellular volume have more time to adjust. We show experimentally that slow salinity changes are less harmful to veligers than rapid ones by isolating capsules from egg-mass gel and exposing them to gradual or abrupt salinity change. The results support the hypothesis that rate of change of salinity is an important determinant of embryo survival and that egg-mass gel retards the rate of salinity change.

Evolutionary relations among vertebrate muscle-type lactate dehydrogenases.

Gene duplication has produced two lactate dehydrogenase (LDH) isozymes, LDH-A and LDH-B, that are found in essentially all vertebrates. On the basis of the biochemical properties of the LDH-A and LDH-B isozymes, it has been suggested that each locus is orthologous among all vertebrates. However, phylogenetic studies have not supported a common evolutionary history among the LDH-A isozymes, particularly when those from lower vertebrates are examined. We present here the sequence of a muscle-type LDH from Fundulus heteroclitus, a teleost fish for which the LDH-B sequence has been determined and shown to be unrelated phylogenetically to tetrapod LDH-A isozymes. Although the sequence of the teleost muscle LDH shares certain features with the LDH-A of tetrapods, phylogenetic analyses do not support an orthologous relation among the LDH-A isozymes of teleost fish and tetrapod vertebrates.

Sequence analysis of teleost retina-specific lactate dehydrogenase C: evolutionary implications for the vertebrate lactate dehydrogenase gene family.

At least two gene duplication events have led to the three lactate dehydrogenase (LDH; EC 1.1.1.27) isozymes (LDH-A, LDH-B, and LDH-C) of chordates. The prevailing model for the evolution of the LDH loci involves duplication of a primordial LDH locus near the origin of vertebrates, giving rise to Ldh-A and Ldh-B. A third locus, designated Ldh-C, is expressed in the spermatocytes of mammals and a single family of birds and in the eye or liver tissues of teleost fishes. Ldh-C might have arisen independently in these taxa as duplications of either Ldh-A or Ldh-B. Several authors have challenged this traditional hypothesis on the basis of amino acid sequence and immunological similarity of the three LDH isozymes. They suggest that the primordial LDH gene was duplicated to form Ldh-C and a locus that later gave rise to Ldh-A and Ldh-B. We have differentiated between these hypotheses by determining the cDNA sequence of the retina-specific LDH-C from a teleost, Fundulus heteroclitus. On the basis of amino acid sequence similarity, we conclude that the LDH-C isozymes in fish and mammals are not orthologous but derive from independent gene duplications. Furthermore, our phylogenetic analyses support previous hypotheses that teleost Ldh-C is derived from a duplication of the Ldh-B locus.