Common fear molecules induce defensive responses in marine prey across trophic levels.

Predator-prey interactions are a key feature of ecosystems and often chemically mediated, whereby individuals detect molecules in their environment that inform whether they should attack or defend. These molecules are largely unidentified, and their discovery is important for determining their ecological role in complex trophic systems. Homarine and trigonelline are two previously identified blue crab (Callinectes sapidus) urinary metabolites that cause mud crabs (Panopeus herbstii) to seek refuge, but it was unknown whether these molecules influence other species within this oyster reef system. In the current study, homarine, trigonelline, and blue crab urine were tested on juvenile oysters (Crassostrea virginica) to ascertain if the same molecules known to alter mud crab behavior also affect juvenile oyster morphology, thus mediating interactions between a generalist predator, a mesopredator, and a basal prey species. Oyster juveniles strengthened their shells in response to blue crab urine and when exposed to homarine and trigonelline in combination, especially at higher concentrations. This study builds upon previous work to pinpoint specific molecules from a generalist predator's urine that induce defensive responses in two marine prey from different taxa and trophic levels, supporting the hypothesis that common fear molecules exist in ecological systems.

Predator signaling of multiple prey on different trophic levels structures trophic cascades.

The capacity of an apex predator to produce nonconsumptive effects (NCEs) in multiple prey trophic levels can create considerable complexity in nonconsumptive cascading interactions, but these effects are poorly studied. We examined such effects in a model food web where the apex predator (blue crabs) releases chemical cues in urine that affect both the intermediate consumer (mud crabs seek shelter) and the basal prey (oysters are induced to grow stronger shells). Shelter availability and predator presence were manipulated in a laboratory experiment to identify patterns in species interactions. Then, experimentally induced and uninduced oysters were planted across high-quality and low-quality habitats with varying levels of shelter availability and habitat heterogeneity to determine the consistency of these patterns in the field. Oyster shell thickening in response to blue crab chemical cues generally protected oysters from mud crab predation in both the laboratory and in field environments that differed in predation intensity, structural complexity, habitat heterogeneity, and predator composition. However, NCEs on the intermediate predator (greater use of refugia) opposed the NCEs on oyster prey in the interior of oyster reefs while still providing survival advantages to basal prey on reef edges and bare substrates. Thus, the combined effects of changing movement patterns of intermediate predators and morphological defenses of basal prey create complex, but predictable, patterns of NCEs across landscapes and ecotones that vary in structural complexity. Generalist predators that feed on multiple trophic levels are ubiquitous, and their potential effects on NCEs propagating simultaneously to different trophic levels must be quantified to understand the role of NCEs in food webs.

Environmental stress gradients regulate the relative importance of predator density- and trait-mediated indirect effects in oyster reef communities.

Predators affect community structure by influencing prey density and traits, but the importance of these effects often is difficult to predict. We measured the strength of blue crab predator effects on mud crab prey consumption of juvenile oysters across a flow gradient that inflicts both physical and sensory stress to determine how the relative importance of top predator density-mediated indirect effects (DMIEs) and trait-mediated indirect effects (TMIEs) change within systems. Overall, TMIEs dominated in relatively benign flow conditions where blue crab predator cues increased oyster survivorship by reducing mud crab-oyster consumption. Blue crab DMIEs became more important in high sensory stress conditions, which impaired mud crab perception of blue crab chemical cues. At high physical stress, the environment benefitted oyster survival by physically constraining mud crabs. Thus, factors that structure communities may be predicted based on an understanding of how physical and sensory performances change across environmental stress gradients.

Morphology and Surface Properties of Roach Water Transport Arrays.

We report on morphological studies of wharf roaches, Ligia exotica, which can passively absorb and transport water through the microscopic protrusions on their legs. We systematically investigated the geometrical variables of the protrusions on each podite of legs to reveal a particularized structural complexity. For the morphological analysis, each podite was split into nine different zones by grouping the protrusions with similar shapes and organization. The protrusions are shown to possess three different types of shapes located on each specific zone of the podite. In addition, the nanoscale surface morphologies of the protrusions on the wharf roach legs were probed by using atomic force microscopy, and the surface properties of the hairy arrays were determined for identifying the localized hydrophobicity distribution. The protrusion surface possessed a nanoscale periodic patterned texture, and both the valley and ridges of a periodic pattern on the protrusion surface exhibited an identical low surface energy. We suggest that the structural morphologies and distinct hydrophobicity of the protrusions can be critical in determining the directional wettability of an entire leg and important for designing a sturdy water transport and passive water-absorbing system without external energy consumption.

Hydrodynamics affect predator controls through physical and sensory stressors.

Predators influence communities through either consuming prey (consumptive effects, CEs) or altering prey traits (non-consumptive effects, NCEs), which has cascading effects on lower trophic levels. CEs are well known to decrease in physically stressful environments, but NCEs may be reduced at physically benign levels that affect the ability of prey to detect and respond to predators (i.e., sensory stress). We investigated the influence of physical and sensory stressors created by spatial and temporal differences in tidal flow on predator controls in a tritrophic system. We estimated mud crab reactive ranges to blue crab NCEs by evaluating mud crab CEs on juvenile oysters at different distances away from caged blue crabs across flow conditions. Mud crab reactive ranges were large at lower physical and sensory stress levels and blue crabs had a positive cascading effect on oyster survival. Blue crab NCEs were not important at higher flow conditions. Oyster survival was a complicated function of both types of stressors. Physical stress (i.e., current speed) had a positive effect on oyster survival by physically limiting mud crab CEs at high current speeds. Sensory stress (i.e., turbulence) interfered with the propagation of blue crab chemical cues used by mud crabs for predator detection, which removed blue crab NCEs. Mud crab CEs increased as a result and had a negative effect on oyster survival in turbulent conditions. Thus, environmental properties, such as fluid flow, can inflict physical and sensory stressors that have distinct effects on basal prey performance through impacts on different predator effects.

Chemical encoding of risk perception and predator detection among estuarine invertebrates.

An effective strategy for prey to survive in habitats rich in predators is to avoid being noticed. Thus, prey are under selection pressure to recognize predators and adjust their behavior, which can impact numerous community-wide interactions. Many animals in murky and turbulent aquatic environments rely on waterborne chemical cues. Previous research showed that the mud crab, Panopeus herbstii, recognizes the predatory blue crab, Callinectus sapidus, via a cue in blue crab urine. This cue is strongest if blue crabs recently preyed upon mud crabs. Subsequently, mud crabs suppress their foraging activity, reducing predation by blue crabs. Using NMR spectroscopy- and mass spectrometry-based metabolomics, chemical variation in urine from blue crabs fed different diets was related to prey behavior. We identified the urinary metabolites trigonelline and homarine as components of the cue that mud crabs use to detect blue crabs, with concentrations of each metabolite dependent on the blue crab's diet. At concentrations found naturally in blue crab urine, trigonelline and homarine, alone as well as in a mixture, alerted mud crabs to the presence of blue crabs, leading to decreased foraging by mud crabs. Risk perception by waterborne cues has been widely observed by ecologists, but the molecular nature of these cues has not been previously identified. Metabolomics provides an opportunity to study waterborne cues where other approaches have historically failed, advancing our understanding of the chemical nature of a wide range of ecological interactions.

Biotic structure indirectly affects associated prey in a predator-specific manner via changes in the sensory environment.

Indirect effects, which can be either positive or negative, may be important in areas containing biotic structure, because such structure can provide refuge and habitat, produce additional sensory cues that may attract predators, and modify the sensory landscape in which predator-prey interactions occur. To determine the indirect effects of biotic structure on prey populations, we assessed predation on patches of hard clams (Mercenaria mercenaria) by large odor-mediated blue crab (Callinectes sapidus) and knobbed whelk (Busycon carica) predators at 0, 5, and 10 m from oyster reefs in intertidal salt marshes. Oyster reefs had an overall indirect negative effect on hard clams, with higher predation rates closer to the reef than farther away. Predator-specific patterns of predation showed that blue crabs consumed more clams very close to the reef, whereas whelks consumed more clams at intermediate distances. Laboratory flume experiments suggest that the oyster reef structure creates turbulence that diminishes predator foraging efficiency, particularly in rapidly mobile predators such as blue crabs, but that oyster reef chemicals ameliorate the negative impact of turbulence on foraging success for both predators. Changes in the sensory landscape, in combination with predator perceptual ability, will determine the positive and/or negative impacts of biotic structure on associated prey. Gaining an understanding of the context specificity of positive and negative sensory effects of biotic structure provides insights that are important for developing a predictive framework to assess the magnitude and distribution of indirect interactions in natural communities.

Predator biomass determines the magnitude of non-consumptive effects (NCEs) in both laboratory and field environments.

Predator body size often indicates predation risk, but its significance in non-consumptive effects (NCEs) and predator risk assessment has been largely understudied. Although studies often recognize that predator body size can cause differing cascading effects, few directly examine prey foraging behavior in response to individual predator sizes or investigate how predator size is discerned. These mechanisms are important since perception of the risk imposed by predators dictates behavioral responses to predators and subsequent NCEs. Here, we evaluate the role of predator body size and biomass on risk assessment and the magnitude of NCEs by investigating mud crab foraging behavior and oyster survival in response to differing biomasses of blue crab predators using both laboratory and field methods. Cues from high predator biomass treatments including large blue crab predators and multiple small blue crab predators decreased mud crab foraging and increased oyster survival, whereas mud crab foraging in response to a single small blue crab did not differ from controls. Mud crabs also increased refuge use in the presence of large and multiple small, but not single small, blue crab predators. Thus, both predator biomass and aggregation patterns may affect the expression of NCEs. Understanding the impact of predator biomass may therefore be necessary to successfully predict the role of NCEs in shaping community dynamics. Further, the results of our laboratory experiments were consistent with observed NCEs in the field, suggesting that data from mesocosm environments can provide insight into field situations where flow and turbulence levels are moderate.

Getting ahead: context-dependent responses to odorant filaments drive along-stream progress during odor tracking in blue crabs.

The chemosensory signal structure governing the upstream progress of blue crabs to an odorant source was examined. We used a three-dimensional laser-induced fluorescence system to collect chemical concentration data simultaneously with behavior observations of actively tracking blue crabs (Callinectes sapidus) in a variety of plume types. This allowed us to directly link chemical signal properties at the antennules and legs to subsequent upstream motion while altering the spatial and temporal intermittency characteristics of the sensory field. Our results suggest that odorant stimuli elicit responses in a binary fashion by causing upstream motion, provided the concentration at the antennules exceeds a specific threshold. In particular, we observed a significant association between crab velocity changes and odorant spike encounters defined using a threshold that is scaled to the mean of the instantaneous maximum concentration. Thresholds were different for each crab, indicating a context-sensitive response to signal dynamics. Our data also indicate that high frequency of odorant spike encounters terminate upstream movement. Further, the data provide evidence that the previous state of the crab and prior stimulus history influence the behavioral response (i.e. the response is context dependent). Two examples are: (1) crabs receiving prior odorant spikes attained elevated velocity more quickly in response to subsequent spikes; and (2) prior acceleration or deceleration of the crab influenced the response time period to a particular odorant spike. Finally, information from both leg and antennule chemosensors interact, suggesting parallel processing of odorant spike properties during navigation.

Staying the course: chemical signal spatial properties and concentration mediate cross-stream motion in turbulent plumes.

This study examined the role of broadly distributed sensor populations in chemosensory searching, especially cross-stream heading adjustment. We used three-dimensional laser-induced fluorescence to collect chemical concentration data simultaneously with behavior observations of actively tracking blue crabs (Callinectes sapidus). Our analysis indicates that the spatial distribution of the odorant concentration field is necessary and sufficient to mediate correct cross-stream motion, although concentration provides information that supplements that obtained from the spatial distribution. Crab movement is continually adjusted to maintain an upstream heading, with corrections toward the source modulated only in the presence of chemical cues. Crabs detect and respond to shifts in the position of the center-of-mass (COM) of the odorant concentration distribution as small as 5% of the leg span, which corresponds to ∼0.8-0.9 cm. The reaction time after a 5% threshold shift in the position of the COM is in the range of 2-4 s. Data also indicate that these steering responses are dependent on stimulus history or other characteristics of the plume, with crabs taking longer to respond in conditions with large-scale spatial meanders. Although cross-stream motion is determined by chemical signal inputs to receptors on the walking legs, crabs do make rotational movements in response to chemical signals impinging on the antennules. These rotational movements do not affect the direction of travel, but rather, determine the crab's body angle with respect to the flow. Interestingly, these body angles seem to represent a compromise between reducing drag and obtaining better chemical signal information, and this trade-off is resolved differently under different plume conditions.

Hydrodynamic sensory stressors produce nonlinear predation patterns.

Predators often have large effects on community structure, but these effects can be minimized in habitats subjected to intense physical stress. For example, predators exert large effects on rocky intertidal communities on wave-protected shores but are usually absent from wave-swept shores where hydrodynamic forces prevent them from foraging effectively. The physical environment also can affect predation levels when stressors are not severe enough to be physically risky. In these situations, environmental conditions may constrain a predator's ability to locate prey and alleviate predation pressure. Yet, stress models of community structure have rarely considered the implications of such sensory or behavioral stressors, particularly when the sensory abilities of both predators and prey are affected by the same types of environmental conditions. Ecologists may classify certain environmental conditions as refuges if they impede predator foraging, but these conditions may not actually decrease predation levels if they simultaneously increase prey vulnerability to consumers. Using blue crabs (Callinectes sapidus) and hard clams (Mercenaria mercenaria) as a model system, we investigated the relationship between predation intensity and environmental stress in the form of hydrodynamics (i.e., flow velocity and turbulence). Blue crabs and hard clams are less responsive to each other in faster, more turbulent flows, but studies exploring how flow modulates the outcomes of crab-clam interactions in the field are lacking. We manipulated turbulence within field sites and compared predation levels within and between sites that differed in flow velocity and turbulence. Our results suggest that blue crabs are most effective foragers in flows with intermediate velocities and turbulence levels. Although these conditions are not ideal for blue crabs, lab studies indicate that they also compromise the ability of clams to detect and react to approaching crabs and, thereby, increase clam vulnerability to predators. Our results suggest that environmental stresses on perception (sensory stressors) may not cause a steady decay in predation rates when they simultaneously affect the behaviors of both predators and prey. Moreover, the relative contribution of lethal vs. nonlethal predator effects in communities also may be influenced by environmental forces that enhance the predator-avoidance abilities of prey or the foraging efficiency of predators.

Alteration of sensory abilities regulates the spatial scale of nonlethal predator effects.

Many studies have shown that nonlethal predator effects such as trait-mediated interactions (TMIs) can have significant impacts on the structure and function of communities, but the role that environmental conditions play in modulating the scale and magnitude of these effects has not been carefully investigated. TMIs occur when prey exhibit behavioral or physiological responses to predators and may be more prevalent when abiotic conditions increase prey reactions to consumers. The purpose of this study was to determine if turbulence would alter the distance over which prey in aquatic systems respond to chemical cues emitted by predators in nature, thus changing the scales over which nonlethal predator effects occur. Using hard clams and blue crabs as a model predator-prey system, we investigated the effects of turbulence on clam reactive distance to predatory blue crabs in the field. Results suggest that turbulence diminishes clam reactions to predators and that the environmental context must be considered when predicting the extent of indirect predator effects in natural systems.

The olfactory pathway mediates sheltering behavior of Caribbean spiny lobsters, Panulirus argus, to conspecific urine signals.

The "noses" of diverse taxa are organized into different subsystems whose functions are often not well understood. The "nose" of decapod crustaceans is organized into two parallel pathways that originate in different populations of antennular sensilla and project to specific neuropils in the brain-the aesthetasc/olfactory lobe pathway and the non-aesthetasc/lateral antennular neuropil pathway. In this study, we investigated the role of these pathways in mediating shelter selection of Caribbean spiny lobsters, Panulirus argus, in response to conspecific urine signals. We compared the behavior of ablated animals and intact controls. Our results show that control and non-aesthetasc ablated lobsters have a significant overall preference for shelters emanating urine over control shelters. Thus the non-aesthetasc pathway does not play a critical role in shelter selection. In contrast, spiny lobsters with aesthetascs ablated did not show a preference for either shelter, suggesting that the aesthetasc/olfactory pathway is important for processing social odors. Our results show a difference in the function of these dual chemosensory pathways in responding to social cues, with the aesthetasc/olfactory lobe pathway playing a major role. We discuss our results in the context of why the noses of many animals contain multiple parallel chemosensory systems.

The prevalence and implications of copepod behavioral responses to oceanographic gradients and biological patchiness.

Several species and developmental stages of calanoid copepods were tested for responses to environmental cues in a laboratory apparatus that mimicked conditions commonly associated with patches of food in the ocean. All species responded to the presence of phytoplankton by feeding. All species responded by increasing proportional residence time in one, but not both, of the treatments defined by gradients of velocity or density. Most species increased swimming speed and frequency of turning in response to the presence of chemical exudates or gradients of velocity. Only one species, Eurytemora affinis, increased proportional time of residence in response to gradients in density of the water. Responses of E. affinis to combined cues did not definitively demonstrate a hierarchical use of different cues as previously observed for Temora longicornis and Acartia tonsa. A simple foraging simulation was developed to assess the applicability in the field of the behavioral results observed in the laboratory. These simulations suggest that observed fine-scale behaviors could lead to copepod aggregations observed in situ. The present study demonstrates that behavioral response to cues associated with fine-scale oceanographic gradients and biological patchiness is functionally important and prevalent among copepods and likely has significant impacts on larger-scale distributional patterns.

Source and specificity of chemical cues mediating shelter preference of Caribbean spiny lobsters (Panulirus argus).

Caribbean spiny lobsters display a diversity of social behaviors, one of the most prevalent of which is gregarious diurnal sheltering. Previous research has demonstrated that shelter selection is chemically mediated, but the source of release and the identity of the aggregation signal are unknown. In this study, we investigated the source and specificity of the aggregation signal in Caribbean spiny lobsters, Panulirus argus. We developed a relatively rapid test of shelter choice in a 5000-l laboratory flume that simulated flow conditions in the spiny lobster's natural environment, and used it to examine the shelter preference of the animals in response to a variety of odorants. We found that both males and females associated preferentially with shelters emanating conspecific urine of either sex, but not with shelters emanating seawater, food odors, or the scent of a predatory octopus. These results demonstrate specificity in the cues mediating sheltering behavior and show that urine is at least one source of the aggregation signal.

Clamming up: environmental forces diminish the perceptive ability of bivalve prey.

The lethal and nonlethal impacts of predators in marine systems are often mediated via reciprocal detection of waterborne chemical signals between consumers and prey. Local flow environments can enhance or impair the chemoreception ability of consumers, but the effect of hydrodynamics on detection of predation risk by prey has not been investigated. Using clams as our model organism, we investigated two specific questions: (1) Can clams decrease their mortality by responding to predators? (2) Do fluid forces affect the ability of clams to detect approaching predators? Previous research has documented a decrease in clam feeding (pumping) in response to a neighboring predator. We determined the benefits of this behavior to survivorship by placing clams in the field with knobbed whelk or blue crab predators caged nearby and compared mortality between these clams and clams near a cage-only control. Significantly more clams survived in areas containing a caged predator, suggesting that predator-induced alterations in feeding reduce clam mortality in the field. We ascertained the effect of fluid forces on clam perception of predators in a laboratory flume by comparing the feeding (pumping) behavior of clams in response to crabs and whelks in flows of 3 and 11 cm/s. Clams pumped significantly less in the presence of predators, but their reaction to blue crabs diminished in the higher velocity flow, while their response to whelks remained constant in both flows. Thus, clam reactive distance to blue crabs was affected by fluid forces, but hydrodynamic effects on clam perceptive distance was predator specific. After predators were removed, clams exposed to whelks took significantly longer to resume feeding than those exposed to blue crabs. Our results suggest that prey perception of predators can be altered by physical forces. Prey detection of predators is the underlying mechanism for trait-mediated indirect interactions (TMIIs), and recent research has documented the importance of TMIIs to community structure. Since physical forces can influence prey perception, the prevalence of TMIIs in communities may, in part, be related to the sensory ability of prey, physical forces in the environment that impact sensory performance, and the type of predator detected.

Hard clams (Mercenaria mercenaria) evaluate predation risk using chemical signals from predators and injured conspecifics.

Hard clams, Mercenaria mercenaria, are sessile, filter-feeding organisms that are heavily preyed upon by blue crabs, which find their clam prey using chemical cues. Clams may evade blue crabs by reducing their pumping (feeding) behavior when a threat is perceived. The purpose of this study was to determine the type of signals that clams use to detect consumers. Clams decreased their pumping time in response to blue crabs and blue crab effluent, but not to crab shells, indicating that chemical signals and not mechanical cues mediated the response of clams to distant predators. Because predator diet can influence prey evaluation of predatory threats, we compared clam responses to blue crabs fed a steady diet of fish, clams, or that were starved prior to the experiment. In addition, we used injured clams as a stimulus because many organisms detect predators by sensing the odor of injured con- or heterospecifics. Clams reduced feeding in response to injured conspecifics and to blue crabs that had recently fed. Clams reacted similarly to fed crabs, regardless of their diet, but did not respond to starved blue crabs. Because blue crabs are generalist predators and the threat posed by these consumers is unrelated to the crab's diet, we should expect clam reactions to blue crabs to be independent of the crab's diet. The failure of clams to react to starved blue crabs likely increases their vulnerability to these consumers, but clam responses to injured conspecifics may constitute a strategy that allows animals to detect an imminent threat when signals emanating from blue crabs are not detectable.

Slow-moving predatory gastropods track prey odors in fast and turbulent flow.

Olfactory searching by aquatic predators is reliant upon the hydrodynamic processes that transport and modify chemical signals. Previous studies indicate that the search behavior of some benthic crustaceans is hindered by rapid water flow and turbulent mixing of prey chemicals, but different sensory strategies employed by other taxa might offset such detrimental effects. Using a laboratory flume, we investigated the odor-tracking behavior of a marine gastropod whelk (Busycon carica) to test the generalization that turbulence interferes with chemically mediated navigation. We exposed individual whelks to turbulent odor plumes in free-stream velocities of 1.5, 5, 10 or 15 cm s(-1), or with one of two obstructions placed upstream of the odor source in an intermediate flow of 5 cm s(-1). Measurements of velocity and stimulus properties confirmed that obstruction treatments increased turbulence intensity and altered the fine-scale structure of downstream odor plumes. In all conditions tested, between 36-63% of test animals successfully located the odor source from 1.5 m downstream with no significant effect of flow treatment. Search behaviors, such as cross-stream meander were reduced at higher flow velocities and in the presence of obstructions, allowing whelks to reach the odor source significantly more quickly than in slower, less turbulent conditions. Our results demonstrate that whelks can respond to chemical information in fast and turbulent flow, and we suggest that these slow-moving predators can forage in hydrodynamic environments where the olfactory abilities of other taxa are limited.

Dual antennular chemosensory pathways can mediate orientation by Caribbean spiny lobsters in naturalistic flow conditions.

Benthic crustaceans rely on chemical stimuli to mediate a diversity of behaviors ranging from food localization and predator avoidance to den selection, conspecific interactions and grooming. To accomplish these tasks, Caribbean spiny lobsters (Panulirus argus) rely on a complex chemosensory system that is organized into two parallel chemosensory pathways originating in diverse populations of antennular sensilla and projecting to distinct neuropils within the brain. Chemosensory neurons associated with aesthetasc sensilla project to the glomerular olfactory lobes (the aesthetasc pathway), whereas those associated with non-aesthetasc sensilla project to the stratified lateral antennular neuropils and the unstructured median antennular neuropil (the non-aesthetasc pathway). Although the pathways differ anatomically, unique roles for each in odor-mediated behaviors have not been established. This study investigates the importance of each pathway for orientation by determining whether aesthetasc or non-aesthetasc sensilla are necessary and sufficient for a lobster to locate the source of a 2 m-distant food odor stimulus in a 5000-liter seawater flume under controlled flow conditions. To assess the importance of each pathway for this task, we selectively ablated specific populations of sensilla on the antennular flagella and compared the searching behavior of ablated animals to that of intact controls. Our results show that either the aesthetasc or the non-aesthetasc pathway alone is sufficient to mediate the behavior and that neither pathway alone is necessary. Under the current experimental conditions, there appears to be a high degree of functional overlap between the pathways for food localization behavior.

Effects of odor flux and pulse rate on chemosensory tracking in turbulent odor plumes by the blue crab, Callinectes sapidus.

The ability of animals to track through chemical plumes is often related to properties of evanescent odor bursts and to small-scale mixing process that determine burst properties. However, odor plumes contain variation over a range of scales, and little is known about how variation in the properties of the odor signal on the scale of one to several seconds affects foraging performance. We examined how flux and pulse rate interact to modulate the search behavior of blue crabs, Callinectes sapidus, locating odor sources in controlled flume flows. Experimental treatments consisted of continuous plumes and plumes with discrete odor pulses at intervals of 2.5 s and 4 s at two fluxes. Crabs experienced diminished search success and reduced search efficiency as flux decreased and the inter-pulse interval lengthened. There often were significant interactions between flux and pulse length, and neither property completely determined search behavior. Thus, over the time span of several seconds, the blue crab chemosensory system is not a simple flux detector. The sensitivity of blue crabs to inter-pulse intervals in the range of several seconds indicates that larger-scale mixing processes, which create odor variation on comparable scales, may exert a significant impact on foraging success in nature.