Et tu, Vibrio cholerae? Kin-cannibalism and a bacterial secretion system.

Type VI secretion systems are molecular syringes used by Gram-negative bacteria to kill heterospecific (non-kin) niche competitors. In this issue of Cell, Mashruwala et al. show that colonies of the pathogen Vibrio cholera can also exhibit T6SS-mediated cell killing of kin cells and that this process benefits emerging resistant mutants, thereby increasing genetic diversity.

Divergent combinations of cis-regulatory elements control the evolution of phenotypic plasticity.

The widespread occurrence of phenotypic plasticity across all domains of life demonstrates its evolutionary significance. However, how plasticity itself evolves and how it contributes to evolution is poorly understood. Here, we investigate the predatory nematode Pristionchus pacificus with its feeding structure plasticity using recombinant-inbred-line and quantitative-trait-locus (QTL) analyses between natural isolates. We show that a single QTL at a core developmental gene controls the expression of the cannibalistic morph. This QTL is composed of several cis-regulatory elements. Through CRISPR/Cas-9 engineering, we identify copy number variation of potential transcription factor binding sites that interacts with a single intronic nucleotide polymorphism. Another intronic element eliminates gene expression altogether, mimicking knockouts of the locus. Comparisons of additional isolates further support the rapid evolution of these cis-regulatory elements. Finally, an independent QTL study reveals evidence for parallel evolution at the same locus. Thus, combinations of cis-regulatory elements shape plastic trait expression and control nematode cannibalism.

Hyperactive Rac stimulates cannibalism of living target cells and enhances CAR-M-mediated cancer cell killing.

The 21kD GTPase Rac is an evolutionarily ancient regulator of cell shape and behavior. Rac2 is predominantly expressed in hematopoietic cells where it is essential for survival and motility. The hyperactivating mutation Rac2E62K also causes human immunodeficiency, although the mechanism remains unexplained. Here, we report that in Drosophila, hyperactivating Rac stimulates ovarian cells to cannibalize neighboring cells, destroying the tissue. We then show that hyperactive Rac2E62K stimulates human HL60-derived macrophage-like cells to engulf and kill living T cell leukemia cells. Primary mouse Rac2+/E62K bone-marrow-derived macrophages also cannibalize primary Rac2+/E62K T cells due to a combination of macrophage hyperactivity and T cell hypersensitivity to engulfment. Additionally, Rac2+/E62K macrophages non-autonomously stimulate wild-type macrophages to engulf T cells. Rac2E62K also enhances engulfment of target cancer cells by chimeric antigen receptor-expressing macrophages (CAR-M) in a CAR-dependent manner. We propose that Rac-mediated cell cannibalism may contribute to Rac2+/E62K human immunodeficiency and enhance CAR-M cancer immunotherapy.

Ultimate sacrifice or opportunistic foraging?: The selective forces behind sexual cannibalism: The selective forces behind sexual cannibalism.

The discovery of rare cases of males cannibalising females sheds new light on the selective forces behind sexual cannibalism in the animal kingdom.

Extra Nestlings That Are Condemned to Die Increase Reproductive Success in Hoopoes.

AbstractThe adaptive value of routinely laying more eggs than can be successfully fledged has intrigued evolutionary biologists for decades. Extra eggs could, for instance, be adaptive as insurance against hatching failures. Moreover, because recent literature demonstrates that sibling cannibalism is frequent in the Eurasian hoopoe (Upupa epops), producing extra offspring that may be cannibalized by older siblings might also be adaptive in birds. Here, directed to explore this possibility in hoopoes, we performed a food supplementation experiment during the laying period and a clutch size manipulation during the hatching stage. We found that females with the food supplement laid on average one more egg than control females and that the addition of a close-to-hatch egg at the end of the hatching period increased the intensity of sibling cannibalism and enhanced fledging success in hoopoe nests. Because none of the extra nestlings from the experimental extra eggs survived until fledging, these results strongly suggest that hoopoes obtain fitness advantages by using temporarily abundant resources to produce additional nestlings that will be cannibalized. These results therefore suppose the first experimental demonstration of the nutritive adaptive function of laying extra eggs in vertebrates with parental care.

Impact of pre-copulatory sexual cannibalism on genetic diversity and efficacy of selection.

Factors that increase reproductive variance among individuals act to reduce effective population size (Ne), which accelerates the loss of genetic diversity and decreases the efficacy of purifying selection. These factors include sexual cannibalism, offspring investment and mating system. Pre-copulatory sexual cannibalism, where the female consumes the male prior to mating, exacerbates this effect. We performed comparative transcriptomics in two spider species, the cannibalistic Trechaleoides biocellata and the non-cannibalistic T. keyserlingi, to generate genomic evidence to support these predictions. First, we estimated heterozygosity and found that genetic diversity is relatively lower in the cannibalistic species. Second, we calculated dN/dS ratios as a measure of purifying selection; a higher dN/dS ratio indicated relaxed purifying selection in the cannibalistic species. These results are consistent with the hypothesis that sexual cannibalism impacts operational sex ratio and demographic processes, which interact with evolutionary forces to shape the genetic structure of populations. However, other factors such as the mating system and life-history traits contribute to shaping Ne. Comparative analyses across multiple contrasting species pairs would be required to disentangle these effects. Our study highlights that extreme behaviours such as pre-copulatory cannibalism may have profound eco-evolutionary effects.

A Theoretical Comparison of Alternative Male Mating Strategies in Cephalopods and Fishes.

We used computer simulations of growth, mating and death of cephalopods and fishes to explore the effect of different life-history strategies on the relative prevalence of alternative male mating strategies. Specifically, we investigated the consequences of single or multiple matings per lifetime, mating strategy switching, cannibalism, resource stochasticity, and altruism towards relatives. We found that a combination of single (semelparous) matings, cannibalism and an absence of mating strategy changes in one lifetime led to a more strictly partitioned parameter space, with a reduced region where the two mating strategies co-exist in similar numbers. Explicitly including Hamilton's rule in simulations of the social system of a Cichlid led to an increase of dominant males, at the expense of both sneakers and dwarf males ("super-sneakers"). Our predictions provide general bounds on the viable ratios of alternative male mating strategies with different life-histories, and under possibly rapidly changing ecological situations.

The evolution of targeted cannibalism and cannibal-induced defenses in invasive populations of cane toads.

Biotic conflict can create evolutionary arms races, in which innovation in one group increases selective pressure on another, such that organisms must constantly adapt to maintain the same level of fitness. In some cases, this process is driven by conflict among members of the same species. Intraspecific conflict can be an especially important selective force in high-density invasive populations, which may favor the evolution of strategies for outcompeting or eliminating conspecifics. Cannibalism is one such strategy; by killing and consuming their intraspecific competitors, cannibals enhance their own performance. Cannibalistic behaviors may therefore be favored in invasive populations. Here, we show that cane toad tadpoles (Rhinella marina) from invasive Australian populations have evolved an increased propensity to cannibalize younger conspecifics as well as a unique adaptation to cannibalism-a strong attraction to vulnerable hatchlings-that is absent in the native range. In response, vulnerable conspecifics from invasive populations have evolved both stronger constitutive defenses and greater cannibal-induced plastic responses than their native range counterparts (i.e., rapid prefeeding development and inducible developmental acceleration). These inducible defenses are costly, incurring performance reductions during the subsequent life stage, explaining why plasticity is limited in native populations where hatchlings are not targeted by cannibalistic tadpoles. These results demonstrate the importance of intraspecific conflict in driving rapid evolution, highlight how plasticity can facilitate adaptation following shifts in selective pressure, and show that evolutionary processes can produce mechanisms that regulate invasive populations.

Size-Dependent Intraguild Predation, Cannibalism, and Resource Allocation Determine the Outcome of Species Coexistence.

AbstractIntraguild predation (IGP), a system in which species compete for resources and prey on each other, is more common than existing theory predicts. In theory, an IG predator and its prey can coexist if the IG predator is a weaker competitor for a shared resource and the predator directly benefits from consuming the prey. However, many species that are IG predators also consume members of their own species (cannibalism). Here, we ask whether cannibalism can help resolve the paradox of IGP systems. Our approach differs from previous work on IGP and cannibalism by explicitly considering the size dependence of predatory interactions and how the benefits of predation are allocated to survival, growth, and fecundity of the predator or cannibal. Our results show that cannibalism facilitates coexistence under conditions that are opposite of those predicted by standard IGP theory: species can coexist when the cannibal is a better competitor on the shared resources, directly benefits little from consuming conspecifics, and allocates resources from predation more toward growth and fecundity over survival. Because the effects of IGP and cannibalism are opposite, when an IGP predator is also a cannibal, coexistence between the IGP predator and its prey is not possible and instead depends on the operation of other coexistence mechanisms (e.g., resource partitioning). These results point to the importance of understanding the relative rates of IGP and cannibalism as well as the resource allocation strategy of the IG predator in determining the likelihood of species coexistence.

Trait-Specific Indirect Effects Underlie Variation in the Response of Spiders to Cannibalistic Social Partners.

AbstractIn cannibalistic species, selection to avoid conspecifics may stem from the need to avoid being eaten or to avoid competition. Individuals may thus use conspecific cues to modulate their behavior to such threats. Yet the nature of variation for such cues remains elusive. Here, we use a half-sib/full-sib design to evaluate the contribution of (indirect) genetic or environmental effects to the behavioral response of the cannibalistic wolf spider Lycosa fasciiventris (Dufour, 1835) toward conspecific cues. Spiders showed variation in relative occupancy time, activity, and velocity on patches with or without conspecific cues, but direct genetic variance was found only for occupancy time. These three traits were correlated and could be lumped in a principal component: spiders spending more time in patches with conspecific cues moved less and more slowly in those areas. Genetic and/or environmental components of carapace width and weight loss in the social partner, which may reflect the quality and/or quantity of cues produced, were significantly correlated with this principal component, with larger partners causing focal individuals to move more slowly. Therefore, environmental and genetic trait variation in social partners may maintain trait diversity in focal individuals, even in the absence of direct genetic variation.

Early Filial Cannibalism in Fish Revisited: Endocrinological Constraint, Costs of Parental Care, and Mating Possibility.

AbstractOffspring desertion by parents generally occurs at an early stage of parental care, which is thought to minimize the costs of parental care prior to desertion. This study investigated the effects of endocrinological constraints on early total filial cannibalism by male Rhabdoblennius nitidus in the field, a paternal brooding blennid fish with androgen-dependent brood cycling. In brood reduction experiments, cannibal males showed low levels of plasma 11-ketotestosterone (11-KT) relative to noncannibals and also similar levels of 11-KT to males in the parental care phase. Since 11-KT regulates male courtship intensity, males with decreased courtship activity would exhibit total filial cannibalism. However, there is a possibility that a transient increase in 11-KT levels at the early stage of parental care delays total filial cannibalism. In contrast, total filial cannibalism could occur before a decline to the lowest 11-KT levels, at which point males might still be able to exhibit courtships, possibly to reduce the costs of parental care. To understand how much and when caregiving males exhibit mating and parental care behaviors, it is important to consider not only the presence of endocrinological constraints but also its intensity and flexibility.

Filial Cannibalism Leads to Chronic Nest Failure of Eastern Hellbender Salamanders (Cryptobranchus alleganiensis).

AbstractIn species that provide parental care, parents will sometimes cannibalize their own young (i.e., filial cannibalism). Here, we quantified the frequency of whole-clutch filial cannibalism in a species of giant salamander (eastern hellbender; Cryptobranchus alleganiensis) that has experienced precipitous population declines with unknown causes. We used underwater artificial nesting shelters deployed across a gradient of upstream forest cover to assess the fates of 182 nests at 10 sites over 8 years. We found strong evidence that nest failure rates increased at sites with low riparian forest cover in the upstream catchment. At several sites, reproductive failure was 100%, mainly due to cannibalism by the caring male. The high incidence of filial cannibalism at degraded sites was not explained by evolutionary hypotheses for filial cannibalism based on poor adult body condition or low reproductive value of small clutches. Instead, larger clutches at degraded sites were most vulnerable to cannibalism. We hypothesize that high frequencies of filial cannibalism of large clutches in areas with low forest cover could be related to changes in water chemistry or siltation that influence parental physiology or that reduce the viability of eggs. Importantly, our results identify chronic nest failure as a possible mechanism contributing to population declines and observed geriatric age structure in this imperiled species.

Diverse strategies that animals use to deter intraspecific predation.

Intraspecific predation is a special case of prey-predator system that is taxonomically widespread. While a heterospecific predator's main goal is to achieve nutrition, intraspecific predation provides conspecifics with other benefits including elimination of competitors for resources and mates. Traditionally intraspecific predation is termed based on the context they occur within, for example egg cannibalism, infanticide and sexual cannibalism. These diverse forms of intraspecific predation and the strategies by which animals are known to deter them are summarized categorically in this review. The diversity of defence strategies animals deploy against conspecific predators is appreciable and raises interesting questions on their evolution for future research.

Costs and benefits of polyandry in a sexually cannibalistic mantis.

Mating with more than one male often provides direct or indirect benefits to female fitness but can also increase the chance of injury and death. Costs of mating are expected to increase linearly with increasing mating number. But how such costs interact with benefits to determine the net payoff of mating multiply is not well understood. Using the highly cannibalistic Springbok mantis, Miomantis caffra, a species where females are stabbed in the abdomen by males during violent premating struggles that males initiate to avoid being cannibalized, we took an experimental approach to assess the economics of polyandry under the risk of external, male-inflicted injury. We predicted that females that mate multiply would be more likely to show abdominal injuries, have higher prereproductive mortality, produce fewer offspring and be more likely to engage in pre-mating cannibalism to avoid unwanted matings. In line with our predictions, we found that the likelihood of abdominal injury was highest among females that mated at least once, and prereproductive death was highest among females that mated twice or three times. Virgin females completely avoided these costs and produced some offspring parthenogenetically but not enough to provide a net benefit. Although mating was better than not mating, there was no singularly optimal mating number: females that mated once and three times produced similarly high numbers of offspring from the first ootheca, which resulted in an intermediate trough in offspring production at two matings. We also found little evidence that cannibalism was deployed as a mate-avoidance strategy: females consistently attacked and consumed males regardless of how many times they mated or how long they were housed with males. Our results suggest the possibility of two distinct mating strategies in M. caffra, where females either mate at a lower frequency to minimize costs or at a higher frequency to maximize benefits. We discuss possible explanations for this bimodal pattern in offspring production.

Viral transmission and infection prevalence in a cannibalistic host-pathogen system.

Cannibalism, while prevalent in the natural world, is often viewed as detrimental to a cannibal's health, especially when they consume pathogen-infected conspecifics. The argument stems from the idea that cannibalizing infected individuals increases the chance of coming into contact with a pathogen and subsequently becoming infected. Using an insect pest, the fall armyworm (Spodoptera frugiperda), that readily cannibalizes at the larval stage and its lethal pathogen, we experimentally examined how cannibalism affects viral transmission at both an individual and population level. Prior to death, the pathogen in the system stops the larval host from growing, resulting in infected individuals being smaller than healthy individuals. This leads to size-structured cannibalism of infected individuals with the larger healthy larvae consuming the smaller infected larvae, which is commonly observed. At the individual level, we show that the probability of cannibalism is relatively high for both infected and uninfected individuals especially when the cannibal is larger than the victim. However, the probability of the cannibal becoming infected given that a pathogen-infected individual has been cannibalized is relatively low. On a population level, when cannibalism is allowed to occur transmission rates decline. Additionally, by cannibalizing infected larvae, cannibals lower the infection risk for non-cannibals. Thus, cannibalism can decrease infection prevalence and, therefore, may not be as deleterious as once thought. Under certain circumstances, cannibalizing infected individuals, from the uninfected host's perspective, may even be advantageous, as one obtains a meal and decreases competition for resources with little chance of becoming infected.

Cannibalistic enemy-pest model: effect of additional food and harvesting.

Biological control using natural enemies with additional food resources is one of the most adopted and ecofriendly pest control techniques. Moreover, additional food is also provided to natural enemies to divert them from cannibalism. In the present work, using the theory of dynamical system, we discuss the dynamics of a cannibalistic predator prey model in the presence of different harvesting schemes in prey (pest) population and provision of additional food to predators (natural enemies). A detailed mathematical analysis and numerical evaluations have been presented to discuss the pest free state, coexistence of species, stability, occurrence of different bifurcations (saddle-node, transcritical, Hopf, Bogdanov-Takens) and the impact of additional food and harvesting schemes on the dynamics of the system. It has been obtained that the multiple coexisting equilibria and their stability depend on the additional food (quality and quantity) and harvesting rates. Interestingly, we also observe that the pest population density decreases immediately even when small amount of harvesting is implemented. Also the eradication of pest population (stable pest free state) could be achieved via variation in the additional food and implemented harvesting schemes. The individual effects of harvesting parameters on the pest density suggest that the linear harvesting scheme is more effective to control the pest population rather than constant and nonlinear harvesting schemes. In the context of biological control programs, the present theoretical work suggests different threshold values of implemented harvesting and appropriate choices of additional food to be supplied for pest eradication.

Spawning ecology of the Saint Paul's Gregory Stegastes sanctipauli, a damselfish endemic to the remote St Peter and St Paul's Archipelago.

We studied the spawning ecology of Stegastes sanctipauli, a damselfish endemic to the small and remote St Peter and St Paul's Archipelago (Brazil), aiming to determine the main correlates of reproductive success, as inferred from rates of egg losses along a spawning season. New clutches appeared mostly in the last quarter moon, when egg predation intensified. Predation by heterospecifics, particularly Halichoeres radiatus (Linnaeus, 1758), accounted for most egg losses, while filial cannibalism showed negligible influence. Predation pressure was highest for larger nests with higher initial density of eggs. Hatching of eggs occurred close to the new moon (i.e., ~5 days after spawning). Most nests occur in shallow reefs (<16 m depth), where main food resources for adults (turf algae) are concentrated, but abundance of egg predators, particularly H. radiatus, is also highest. This latter result suggests that the spawning activity of S. sanctipauli is influenced by the trade-offs between food accessibility by adults and egg predation risk. The patterns recorded here are similar to those known for other tropical damselfish species and may reflect strategies for attenuating predation pressure over new fry and eggs.

Cannibalism-overview and medicolegal issues.

Cannibalism, the consumption of another by an individual of the same species, is a widespread practice amongst many animal groups. Human cannibalism or anthropophagy, however, is less common but has been found in many diverse groups ranging from hominids to Crusaders and soldiers in World War II. Although the existence of human cannibalism has been vigorously debated in recent times, it seems clear that well-described cases have occurred. The motivation for consuming human tissues may be (1) nutritional, (2) ritual and (3) pathological. A case of alleged cannibalism involving one of the victims of the so-called Snowtown serial killings in South Australia, Australia, is reported with an analysis of the history and features of cannibalism. Forensic problems may occur in accurately identifying remains that have been cannibalized; however, if ritualistic, serial and/or sadistic homicides are encountered, cannibalism should be considered, particularly if body parts are missing.

Delayed early life effects in the threespine stickleback.

Early life conditions can have a decisive influence on viability later in life. However, the influence of embryo density within a nest or body cavity on subsequent viability has received little attention within an ecological setting. This is surprising given that embryos often compete for limited resources, such as nutrients and oxygen, and this could influence their viability later in life through carry-over and compensatory effects. We show that the density of fertilized eggs within the nests of threespine stickleback males (Gasterosteus aculeatus) influences their viability after hatching. Embryos from larger broods hatch earlier and at a smaller size than those from smaller broods, which reduces their survival until the age of four weeks. This indicates a trade-off between the number and viability of offspring that males can raise to the hatching stage, which could explain the high incidence of partial egg cannibalism in nest-brooding fishes-as a strategy to improve the survival of remaining offspring. These results highlight the importance of considering conditions at the embryonic stage when evaluating the impact of early life conditions on viability and the adaptive value of reproductive decisions.

Drosophila melanogaster cloak their eggs with pheromones, which prevents cannibalism.

Oviparous animals across many taxa have evolved diverse strategies that deter egg predation, providing valuable tests of how natural selection mitigates direct fitness loss. Communal egg laying in nonsocial species minimizes egg predation. However, in cannibalistic species, this very behavior facilitates egg predation by conspecifics (cannibalism). Similarly, toxins and aposematic signaling that deter egg predators are often inefficient against resistant conspecifics. Egg cannibalism can be adaptive, wherein cannibals may benefit through reduced competition and added nutrition, but since it reduces Darwinian fitness, the evolution of anticannibalistic strategies is rife. However, such strategies are likely to be nontoxic because deploying toxins against related individuals would reduce inclusive fitness. Here, we report how D. melanogaster use specific hydrocarbons to chemically mask their eggs from cannibal larvae. Using an integrative approach combining behavioral, sensory, and mass spectrometry methods, we demonstrate that maternally provisioned pheromone 7,11-heptacosadiene (7,11-HD) in the eggshell's wax layer deters egg cannibalism. Furthermore, we show that 7,11-HD is nontoxic, can mask underlying substrates (for example, yeast) when coated upon them, and its detection requires pickpocket 23 (ppk23) gene function. Finally, using light and electron microscopy, we demonstrate how maternal pheromones leak-proof the egg, consequently concealing it from conspecific larvae. Our data suggest that semiochemicals possibly subserve in deceptive functions across taxa, especially when predators rely on chemical cues to forage, and stimulate further research on deceptive strategies mediated through nonvisual sensory modules. This study thus highlights how integrative approaches can illuminate our understanding on the adaptive significance of deceptive defenses and the mechanisms through which they operate.