Fasting or fear: disentangling the roles of predation risk and food deprivation in the nitrogen metabolism of consumers.

Predators can alter nutrient cycles simply by inducing stress in prey. This stress accelerates prey's protein catabolism, nitrogen waste production, and nitrogen cycling. Yet predators also reduce the feeding rates of their prey, inducing food deprivation that is expected to slow protein catabolism and nitrogen cycling. The physiology of prey under predation risk thus balances the influences of predation risk and food deprivation, and this balance is central to understanding the role of predators in nutrient cycles. We explored the separate and combined effects of predation risk and food deprivation on prey physiology and nutrient cycling by exposing guppies (Poecilia reticulata) to predation risk and food deprivation in a 2 × 2 design. We simulated predation risk using chemical cues from a natural predator of guppies, and we created food deprivation by rationing food availability. We measured guppy response as food consumption, growth, tissue energy density, tissue carbon:nitrogen, and nitrogen (N) excretion and assimilation. We found that N-linked physiological processes (N consumption, assimilation, excretion) were strongly affected by predation risk, independent of food consumption. Guppies excreted substantially less under predation risk than they did under food deprivation or control conditions. These results suggest that predation risk, per se, triggers physiological changes in guppies that increase N retention and decrease N excretion. We suggest that slower N metabolism under predation risk is an adaptive response that minimizes protein loss in the face of predictable, predator-induced food restriction. Notably, N metabolism shares common hormonal control with food seeking behavior, and we speculate that increased N retention is a direct and immediate result of reduced food seeking under predation risk. Contrary to predation-stress-based hypotheses for how predators affect nutrient cycling by prey, our result indicates that even short-term exposure to predators may decelerate, rather than accelerate, the speed of N cycling by suppressing N turnover by prey.

A global database of nitrogen and phosphorus excretion rates of aquatic animals.

Animals can be important in modulating ecosystem-level nutrient cycling, although their importance varies greatly among species and ecosystems. Nutrient cycling rates of individual animals represent valuable data for testing the predictions of important frameworks such as the Metabolic Theory of Ecology (MTE) and ecological stoichiometry (ES). They also represent an important set of functional traits that may reflect both environmental and phylogenetic influences. Over the past two decades, studies of animal-mediated nutrient cycling have increased dramatically, especially in aquatic ecosystems. Here we present a global compilation of aquatic animal nutrient excretion rates. The dataset includes 10,534 observations from freshwater and marine animals of N and/or P excretion rates. These observations represent 491 species, including most aquatic phyla. Coverage varies greatly among phyla and other taxonomic levels. The dataset includes information on animal body size, ambient temperature, taxonomic affiliations, and animal body N:P. This data set was used to test predictions of MTE and ES, as described in Vanni and McIntyre (2016; Ecology DOI: 10.1002/ecy.1582).

Changes in digestive traits and body nutritional composition accommodate a trophic niche shift in Trinidadian guppies.

A trophic niche shift can occur as an adaptive response to environmental change such as altered resource quality, abundance or composition. Alterations in digestive traits such as gut morphology and physiology may enable these niche shifts and affect the persistence of populations and species. Relatively few studies, however, have assessed how niche shifts influence suites of digestive traits through phenotypic plasticity and evolutionary mechanisms, and how these trait changes can subsequently alter the nutrition, fitness and life history of organisms. We investigated how population divergence and plasticity alter the gut physiology of wild Trinidadian guppies (Poecilia reticulata), assessing whether variation in digestive traits correspond with enhanced nutrient assimilation under a pronounced dietary shift. We examined gut enzyme activity, and gut size and mass of wild guppies from both high-predation (HP) and low-predation (LP) habitats when reared in the laboratory and fed on high- or low-quality diets designed to reflect their dietary differences previously found in nature. After 10 weeks on the experimental diets, HP guppies maintained shorter and lighter guts than LP guppies on either diet. Guppies also differed in their digestive enzymatic profiles, more often reflecting nutrient balancing so that increased enzyme expression tended to correspond with more deficient nutrients in the diet. LP guppies had increased somatic phosphorus at the end of the experiment, possibly related to the higher alkaline phosphatase activity in their guts. Our results suggest that differences in gut physiology exist among populations of Trinidadian guppies that may reflect local adaptation to their disparate environments.

Metabolic stoichiometry and the ecology of fear in Trinidadian guppies: consequences for life histories and stream ecosystems.

Consumer-driven nutrient recycling, the release of chemicals as byproducts and excesses of consumer physiology, can alter ecosystems by changing the availability of limiting nutrients at the base of the food web. The mere presence of predators can alter consumer physiology by restricting food intake and inducing stress. Predation risk, then, can influence ecosystem function by modifying the role of prey as nutrient recyclers, yet there are few empirical tests of how predation risk alters nutrient recycling by prey. Here, we present the results of a test for the effects of predation risk on the C and N budgets of Trinidadian guppies (Poecilia reticulata). We reared female guppies for 7 weeks on diets of varying quality, and we compared control individuals to those exposed continuously to chemical cues emitted by a guppy predator, Crenicichla alta. We measured food consumption, growth rate, tissue elemental stoichiometry and N excretion by guppies on all treatments. Guppies strongly reduced food intake in the presence of predator cues; however, cue-exposed guppies assimilated nutrients more efficiently than controls. Specifically, cue-exposed guppies strongly increased N retention efficiency while only moderately increasing C efficiency. Consequently, guppies reared with predator cues excreted 39% less N than control guppies. We suggest that reduced foraging, enhanced nutrient efficiency, and decreased N excretion are adaptive responses to the extrinsic mortality threat posed by guppy predators. The resulting substantial reduction in N excretion by guppies may influence ecosystem function in natural streams by reducing the supply of a limiting nutrient.

The influence of dietary and whole-body nutrient content on the excretion of a vertebrate consumer.

In many contexts, nutrient excretion by consumers can impact ecosystems by altering the availability of limiting nutrients. Variation in nutrient excretion can be predicted by mass balance models, most of which are premised on two key ideas: (1) consumers maintain fixed whole-body nutrient content (i.e., %N and %P), so-called fixed homeostasis; (2) if dietary nutrients are not matched to whole-body nutrients, excesses of any nutrient are released as excretion to maintain fixed homeostasis. Mass balance models thus predict that consumer excretion should be positively correlated with diet nutrients and negatively correlated with whole-body nutrients. Recent meta-analyses and field studies, however, have often failed to find these expected patterns, potentially because of a confounding influence-flexibility in whole-body nutrient content with diet quality (flexible homeostasis). Here, we explore the impact of flexible homeostasis on nutrient excretion by comparing the N and P excretion of four genetically diverged Trinidadian guppy (Poecilia reticulata) populations when reared on diets of variable P content. As predicted by mass balance, P excretion increased on the high-P diet, but, contrary to the notion of fixed homeostasis, guppy whole-body %P also increased on the high-P diet. While there was no overall correlation between excretion nutrients and whole-body nutrients, when the effect of diet on both whole-body and excretion nutrients was included, we detected the expected negative correlation between whole-body N:P and excretion N:P. This last result suggests that mass balance can predict excretion rates within species, but only if dietary effects on whole-body nutrient content are controlled. Flexible homeostasis can obscure patterns predicted by mass balance, creating an imperative to accurately capture an organism's diet quality in predicting its excretion rate.

Population variation in the trophic niche of the Trinidadian guppy from different predation regimes.

Population variation in trophic niche is widespread among organisms and is of increasing interest given its role in both speciation and adaptation to changing environments. Trinidadian guppies (Poecilia reticulata) inhabiting stream reaches with different predation regimes have rapidly evolved divergent life history traits. Here, we investigated the effects of both predation and resource availability on guppy trophic niches by evaluating their gut contents, resource standing stocks, and δ15N and δ13C stable isotopes across five streams during the wet season. We found that guppies from low predation (LP) sites had a consistently higher trophic position and proportion of invertebrates in their guts and assimilate less epilithon than guppies from high predation (HP) sites. Higher trophic position was also associated with lower benthic invertebrate availability. Our results suggest that LP guppies could be more efficient invertebrate consumers, possibly as an evolutionary response to greater intraspecific competition for higher quality food. This may be intensified by seasonality, as wet season conditions can alter resource availability, feeding rates, and the intensity of intraspecific competition. Understanding how guppy diets vary among communities is critical to elucidating the role of niche shifts in mediating the link between environmental change and the evolution of life histories.

Substantiating Appropriate Motion Capture Techniques for the Assessment of Nordic Walking Gait and Posture in Older Adults.

Nordic walking (NW) has become a safe and simple form of exercise in recent years, and in studying this gait pattern, various data collection techniques have been employed, each with positives and negatives. The aim was to determine the effect of NW on older adult gait and posture and to determine optimal use of different data collection systems in both short and long duration analysis. Gait and posture during NW and normal walking were assessed in 17 healthy older adults (age: 69 ± 7.3). Participants performed two trials of 6 Minute Walk Tests (6MWT) (1 with poles (WP) and 1 without poles (NP)) and 6 trials of a 5m walk (3 WP and 3 NP). Motion was recorded using two systems, a 6-sensor accelerometry system and an 8-camera 3-dimensional motion capture system, in order to quantify spatial-temporal, kinematic, and kinetic parameters. With both systems, participants demonstrated increased stride length and double support and decreased gait speed and cadence WP compared to NP (p <0.05). Also, with motion capture, larger single support time was found WP (p <0.05). With 3-D capture, smaller hip power generation and moments of force were found at heel contact and pre-swing as well as smaller knee power absorption at heel contact, pre-swing, and terminal swing WP compared to NP, when assessed over one cycle (p <0.05). Also, WP yielded smaller moments of force at heel contact and terminal swing along with larger moments at mid-stance of a gait cycle (p <0.05). No changes were found for posture. NW seems appropriate for promoting a normal gait pattern in older adults. Three-dimensional motion capture should primarily be used during short duration gait analysis (i.e. single gait cycle), while accelerometry systems should be primarily employed in instances requiring longer duration analysis such as during the 6MWT.

Divergence across diet, time and populations rules out parallel evolution in the gut microbiomes of Trinidadian guppies.

Diverse microbial consortia profoundly influence animal biology, necessitating an understanding of microbiome variation in studies of animal adaptation. Yet, little is known about such variability among fish, in spite of their importance in aquatic ecosystems. The Trinidadian guppy, Poecilia reticulata, is an intriguing candidate to test microbiome-related hypotheses on the drivers and consequences of animal adaptation, given the recent parallel origins of a similar ecotype across streams. To assess the relationships between the microbiome and host adaptation, we used 16S rRNA amplicon sequencing to characterize gut bacteria of two guppy ecotypes with known divergence in diet, life history, physiology and morphology collected from low-predation (LP) and high-predation (HP) habitats in four Trinidadian streams. Guts were populated by several recurring, core bacteria that are related to other fish associates and rarely detected in the environment. Although gut communities of lab-reared guppies differed from those in the wild, microbiome divergence between ecotypes from the same stream was evident under identical rearing conditions, suggesting host genetic divergence can affect associations with gut bacteria. In the field, gut communities varied over time, across streams and between ecotypes in a stream-specific manner. This latter finding, along with PICRUSt predictions of metagenome function, argues against strong parallelism of the gut microbiome in association with LP ecotype evolution. Thus, bacteria cannot be invoked in facilitating the heightened reliance of LP guppies on lower-quality diets. We argue that the macroevolutionary microbiome convergence seen across animals with similar diets may be a signature of secondary microbial shifts arising some time after host-driven adaptation.

Predator-induced phenotypic plasticity in metabolism and rate of growth: rapid adaptation to a novel environment.

Novel environments often impose directional selection for a new phenotypic optimum. Novel environments, however, can also change the distribution of phenotypes exposed to selection by inducing phenotypic plasticity. Plasticity can produce phenotypes that either align with or oppose the direction of selection. When plasticity and selection are parallel, plasticity is considered adaptive because it provides a better pairing between the phenotype and the environment. If the plastic response is incomplete and falls short of producing the optimum phenotype, synergistic selection can lead to genetic divergence and bring the phenotype closer to the optimum. In contrast, non-adaptive plasticity should increase the strength of selection, because phenotypes will be further from the local optimum, requiring antagonistic selection to overcome the phenotype-environment mismatch and facilitate adaptive divergence. We test these ideas by documenting predator-induced plasticity for resting metabolic rate and growth rate in populations of the Trinidadian guppy (Poecilia reticulata) adapted to high and low predation. We find reduced metabolic rates and growth rates when cues from a predator are present during development, a pattern suggestive of adaptive and non-adaptive plasticity, respectively. When we compared populations recently transplanted from a high-predation environment into four streams lacking predators, we found evidence for rapid adaptive evolution both in metabolism and growth rate. We discuss the implications for predicting how traits will respond to selection, depending on the type of plasticity they exhibit.

Fates beyond traits: ecological consequences of human-induced trait change.

Human-induced trait change has been documented in freshwater, marine, and terrestrial ecosystems worldwide. These trait changes are driven by phenotypic plasticity and contemporary evolution. While efforts to manage human-induced trait change are beginning to receive some attention, managing its ecological consequences has received virtually none. Recent work suggests that contemporary trait change can have important effects on the dynamics of populations, communities, and ecosystems. Therefore, trait changes caused by human activity may be shaping ecological dynamics on a global scale. We present evidence for important ecological effects associated with human-induced trait change in a variety of study systems. These effects can occur over large spatial scales and impact system-wide processes such as trophic cascades. Importantly, the magnitude of these effects can be on par with those of traditional ecological drivers such as species presence. However, phenotypic change is not always an agent of ecological change; it can also buffer ecosystems against change. Determining the conditions under which phenotypic change may promote vs prevent ecological change should be a top research priority.