Selection for cuticular melanism reveals immune function and life-history trade-offs in Spodoptera littoralis

Several insect species show an increase in cuticular melanism in response to high densities. In some species, there is evidence that this melanism is correlated with an up-regulation of certain immune system components, particularly phenoloxidase (PO) activity, and with the down-regulation of lysozyme activity, suggesting a trade-off between the two traits. As melanism has a genetic component, we selected both melanic and nonmelanic lines of the phase-polyphenic lepidopteran, Spodoptera littoralis, in order to test for a causative genetic link between melanism, PO activity and lysozyme activity, and to establish if there are any life-history costs associated with the melanic response. We found that, in fact, melanic lines had lower PO activity and higher lysozyme activity than nonmelanic lines, confirming a genetic trade-off between the two immune responses, but also indicating a genetic trade-off between melanism and PO activity. In addition, we found that lines with high PO activity had slower development rates suggesting that investment in PO, rather than in melanism, is costly.

Male morph predicts investment in larval immune function in the dung beetle, Onthophagus taurus

Investment in immunity is costly, so that resource-based trade-offs between immunity and sexually selected ornaments might be expected. The amount of resources that an individual can invest in each trait will be limited by the total resources available to them. It would therefore be informative to investigate how investment in immune function changes during growth or production of the sexual trait as resources are diverted to it. Using the dung beetle, Onthophagus taurus, which displays both sexual and male dimorphism in horn size, we examined changes in one measure of immune function, phenoloxidase (PO) activity, in the hemolymph of larvae prior to and during horn growth. We found that PO levels differed between small- and large-horned males throughout the final instar prior to the point where investment in horn growth was taking place. PO levels in females were intermediate to the 2 male morphs. These differences could not be accounted for by differences in condition, measured as hemolymph protein levels and weight. We suggest that the observed differences might be associated with sex- and morph-specific variation in juvenile hormone levels.

Density-dependent prophylaxis and condition-dependent immune function in Lepidopteran larvae: a multivariate approach

1. The risk of parasitism and infectious disease is expected to increase with population density as a consequence of positive density-dependent transmission rates. Therefore, species that encounter large fluctuations in population density are predicted to exhibit plasticity in their immune system, such that investment in costly immune defences is adjusted to match the probability of exposure to parasites and pathogens (i.e. density-dependent prophylaxis).

Costs of resistance: genetic correlations and potential trade-offs in an insect immune system

Theory predicts that natural selection will erode additive genetic variation in fitness-related traits. However, numerous studies have found considerable heritable variation in traits related to immune function, which should be closely linked to fitness. This could be due to trade-offs maintaining variation in these traits. We used the Egyptian cotton leafworm, Spodoptera littoralis, as a model system to examine the quantitative genetics of insect immune function. We estimated the heritabilities of several different measures of innate immunity and the genetic correlations between these immune traits and a number of life history traits. Our results provide the first evidence for a potential genetic trade-off within the insect immune system, with antibacterial activity (lysozyme-like) exhibiting a significant negative genetic correlation with haemocyte density, which itself is positively genetically correlated with both haemolymph phenoloxidase activity and cuticular melanization. We speculate on a potential trade-off between defence against parasites and predators, mediated by larval colour, and its role in maintaining genetic variation in traits under natural selection.

Heritability of immune function in the caterpillar Spodoptera littoralis

Phenoloxidase (PO) is believed to be a key mediator of immune function in insects and has been implicated both in non-self recognition and in resistance to a variety of parasites and pathogens, including baculoviruses and parasitoids. Using larvae of the Egyptian cotton leafworm, Spodoptera littoralis, we found that despite its apparent importance, haemolymph PO activity varied markedly between individuals, even amongst insects reared under apparently identical conditions. Sib-analysis methods were used to determine whether individuals varied genetically in their PO activity, and hence in one aspect of immune function. The heritability estimate of haemolymph PO activity was high (h 2 = 0.690 +/- 0.069), and PO activity in the haemolymph was strongly correlated with PO activity in both the cuticle and midgut; the sites of entry for most parasites and pathogens. Haemolymph PO activity was also strongly correlated with the degree to which a synthetic parasite (a small piece of nylon monofilament) was encapsulated and melanized (r = 0.622 +/- 0.142), suggesting that the encapsulation response is also heritable. The mechanism maintaining this genetic variation has yet to be elucidated.

Macronutrient balance mediates trade-offs between immune function and life history traits

1. Diet and health are intimately linked and recent studies have found that caloric restriction can affect immune function. However, when given a choice between diets that differ in their macronutrient composition, pathogen-infected individuals can select a diet that improves their survival, suggesting that the nutritional composition of the diet, as well as its calorie content, can play a role in defence against disease. Moreover, as individuals change their diet when infected, it suggests that a diet that is optimal for growth is not optimal for immunity, leading to trade-offs.
2. Currently, our knowledge of the effects of diet on immunity is limited because previous experiments have manipulated either single nutrients or the calorie content of the diet without considering their interactive effects. By simultaneously manipulating both the diet composition (quality) and its caloric density (quantity), in both naive and immune-challenged insects, we asked how do diet quality and quantity influence an individual's ability to mount an immune response? And to what extent are allocation trade-offs driven by quantity- versus quality-based constraints?
3. We restricted individuals to 20 diets varying in their protein and carbohydrate content and used 3D response surfaces to visualize dietary effects on larval growth and immune traits. Our results show that both constitutive and induced immune responses are not limited by the total quantity of nutrients consumed, but rather different traits respond differently to variation in the ratios of macronutrients (diet quality), and peak in different regions of macronutrient space. The preferred dietary composition therefore represents a compromise between the nutritional requirements of growth and immune responses. We also show that a non-pathogenic immune challenge does not affect diet choice, rather immune-challenged insects modify their allocation of nutrients to improve their immune response.
4. Our results indicate that immune traits are affected by the macronutrient content of the diet and that no diet can simultaneously optimize all components of the immune system. To date the emphasis has been on the effects of micronutrients in improving immunity, our findings indicate that this must be widened to include the neglected impact of macronutrients on defence against disease.

Oblique amplitude modulation of dust-acoustic plasma waves

Theoretical and numerical studies are presented of the nonlinear amplitude modulation of dust-acoustic (DA) waves propagating in an unmagnetized three component, weakly-coupled, fully ionized plasma consisting of electrons, positive ions and charged dust particles, considering perturbations oblique to the carrier wave propagation direction. The stability analysis, based on a nonlinear Schrodinger-type equation (NLSE), shows that the wave may become unstable; the stability criteria depend on the angle theta between the modulation and propagation directions. Explicit expressions for the instability rate and threshold have been obtained in terms of the dispersion laws of the system. The possibility and conditions for the existence of different types of localized excitations have also been discussed.

Ion-acoustic waves in a two-electron-temperatute plasma: oblique modulation and envelope excitations

Theoretical and numerical studies are carried out for the nonlinear amplitude modulation of ion-acoustic waves propagating in an unmagnetized, collisionless, three-component plasma composed of inertial positive ions moving in a background of two thermalized electron populations. Perturbations oblique to the carrier wave propagation direction have been considered. The stability analysis, based on a nonlinear Schrodinger-type equation, shows that the wave may become unstable; the stability criteria depend on the angle theta between the modulation and propagation directions. Different types of localized excitations (envelope solitary waves) are shown to exist in qualitative agreement with satellite observations in the magnetosphere.

Electron-acoustic plasma waves: Oblique modulation and envelope solitons

Theoretical and numerical studies are presented of the amplitude modulation of electron-acoustic waves (EAWs) propagating in space plasmas whose constituents are inertial cold electrons, Boltzmann distributed hot electrons, and stationary ions. Perturbations oblique to the carrier EAW propagation direction have been considered. The stability analysis, based on a nonlinear Schrodinger equation, reveals that the EAW may become unstable; the stability criteria depend on the angle theta between the modulation and propagation directions. Different types of localized EA excitations are shown to exist.

Nonlinear modulation of transverse dust lattice waves in complex plasma crystals

The occurrence of the modulational instability in transverse dust lattice waves propagating in a one-dimensional dusty plasma crystal is investigated. The amplitude modulation mechanism, which is related to the intrinsic nonlinearity of the sheath electric field, is shown to destabilize the carrier wave under certain conditions, possibly leading to the formation of localized envelope excitations. Explicit expressions for the instability growth rate and threshold are presented and discussed. (C) 2004 American Institute of Physics.

Modulational instability of dust acoustic waves in dusty plasmas: Modulation obliqueness, background ion nonthermality, and dust charging effects

The oblique modulational instability of dust acoustic (DA) waves in an unmagnetized warm dusty plasma with nonthermal ions, taking into account dust grain charge variation (charging), is investigated. A nonlinear Schrodinger-type equation governing the slow modulation of the wave amplitude is derived. The effects of dust temperature, dust charge variation, ion deviation from Maxwellian equilibrium (nonthermality) and constituent species' concentration on the modulational instability of DA waves are examined. It is found that these parameters modify significantly the oblique modulational instability domain in the k-theta plane. Explicit expressions for the instability rate and threshold have been obtained in terms of the dispersion laws of the system. The possibility and conditions for the existence of different types of localized excitations are also discussed. The findings of this investigation may be useful in understanding the stable electrostatic wave packet acceleration mechanisms close to the Moon, and also enhances our knowledge on the occurrence of instability associated to pickup ions around unmagnetized bodies, such as comets, Mars, and Venus.

Oblique modulation of electrostatic modes and envelope excitations in pair-ion and electron-positron plasmas

The nonlinear amplitude modulation of electrostatic waves propagating in a collisionless two-component plasma consisting of negative and positive species of equal mass and absolute charge is investigated. Pair-ion (e.g., fullerene) and electron-positron (e-p) plasmas (neglecting recombination) are covered by this description. Amplitude perturbation oblique to the direction of propagation of the wave has been considered. Two distinct linear electrostatic modes exist, namely an acoustic lower mode and Langmuir-type optic-type upper one. The behavior of each of these modes is examined from the modulational stability point of view. The stability criteria are investigated, depending on the electrostatic carrier wave number, the angle theta between the modulation and propagation directions, and the positron-to-electron temperature ratio sigma. The analysis shows that modulated electrostatic wavepackets associated to the lower (acoustic) mode are unstable, for small values of carrier wave number k (i.e., for large wavelength lambda) and for finite (small) values of the angle theta (yet stable for higher theta), while those related to the upper (optic-like) mode are stable for large values of the angle theta only, in the same limit, yet nearly for all values of sigma. These results are of relevance in astrophysical contexts (e.g., in pulsar environments), where e-p plasmas are encountered, or in pair fullerene-ion plasmas, in laboratory. (c) 2006 American Institute of Physics.

Dust-acoustic wave modulation in the presence of superthermal ions

A study is presented of the nonlinear self-modulation of low-frequency electrostatic (dust acoustic) waves propagating in a dusty plasma, in the presence of a superthermal ion (and Maxwellian electron) background. A kappa-type superthermal distribution is assumed for the ion component, accounting for an arbitrary deviation from Maxwellian equilibrium, parametrized via a real parameter kappa. The ordinary Maxwellian-background case is recovered for kappa ->infinity. By employing a multiple scales technique, a nonlinear Schrodinger-type equation (NLSE) is derived for the electric potential wave amplitude. Both dispersion and nonlinearity coefficients of the NLSE are explicit functions of the carrier wavenumber and of relevant physical parameters (background species density and temperature, as well as nonthermality, via kappa). The influence of plasma background superthermality on the growth rate of the modulational instability is discussed. The superthermal feature appears to control the occurrence of modulational instability, since the instability window is strongly modified. Localized wavepackets in the form of either bright-or dark-type envelope solitons, modeling envelope pulses or electric potential holes (voids), respectively, may occur. A parametric investigation indicates that the structural characteristics of these envelope excitations (width, amplitude) are affected by superthermality, as well as by relevant plasma parameters (dust concentration, ion temperature).

Nonlinear modulation of ion-acoustic waves in two-electron-temperature plasmas

The amplitude modulation of ion-acoustic waves IS investigated in a plasma consisting of adiabatic warm ions, and two different populations of thermal electrons at different temperatures. The fluid equations are reduced to nonlinear Schrodinger equation by employing a multi-scale perturbation technique. A linear stability analysis for the wave packet amplitude reveals that long wavelengths are always stable, while modulational instability sets in for shorter wavelengths. It is shown that increasing the value of the hot-to-cold electron temperature ratio (mu), for a given value of the hot-to-cold electron density ratio (nu): favors instability. The role of the ion temperature is also discussed. In the limiting case nu = 0 (or nu -> infinity). which correspond(s) to an ordinary (single) electron-ion plasma, the results of previous works are recovered.