Small skeletons show size-specific scaling: an exploration of allometry in the mammalian lumbar spine.

Studies of vertebrate bone biomechanics often focus on skeletal adaptations at upper extremes of body mass, disregarding the importance of skeletal adaptations at lower extremes. Yet mammals are ancestrally small and most modern species have masses under 5 kg, so the evolution of morphology and function at small size should be prioritized for understanding how mammals subsist. We examined allometric scaling of lumbar vertebrae in the small-bodied Philippine endemic rodents known as cloud rats, which vary in mass across two orders of magnitude (15.5 g-2700 g). External vertebral dimensions scale with isometry or positive allometry, likely relating to body size and nuances in quadrupedal posture. In contrast to most mammalian trabecular bone studies, bone volume fraction and trabecular thickness scale with positive allometry and isometry, respectively. It is physiologically impossible for these trends to continue to the upper extremes of mammalian body size, and we demonstrate a fundamental difference in trabecular bone allometry between large- and small-bodied mammals. These findings have important implications for the biomechanical capabilities of mammalian bone at small body size; for the selective pressures that govern skeletal evolution in small mammals; and for the way we define 'small' and 'large' in the context of vertebrate skeletons.

Body mass mediates spatio-temporal responses of mammals to human frequentation across Italian protected areas.

Protected area (PA) networks are a pivotal tool to fight biodiversity loss, yet they often need to balance the mission of nature conservation with the socio-economic need of giving opportunity for outdoor recreation. Recreation in natural areas is important for human health in an urbanized society, but can prompt behavioural modifications in wild animals. Rarely, however, have these responses being studied across multiple PAs and using standardized methods. We deployed a systematic camera trapping protocol at over 200 sites to sample medium and large mammals in four PAs within the European Natura 2000 network to assess their spatio-temporal responses to human frequentation, proximity to towns, amount of open habitat and topographical variables. By applying multi-species and single-species models for the number of diurnal, crepuscular and nocturnal detections and a multi-species model for nocturnality index, we estimated both species-specific- and meta-community-level effects, finding that increased nocturnality appeared the main strategy that the mammal meta-community used to cope with human disturbance. However, responses in the diurnal, crepuscular and nocturnal site use were mediated by species' body mass, with larger species exhibiting avoidance of humans and smaller species more opportunistic behaviours. Our results show the effectiveness of standardized sampling and provide insights for planning the expansion of PA networks as foreseen by the Kunming-Montreal biodiversity agreement.

Parental brain through time: The origin and development of the neural circuit of mammalian parenting.

This review consolidates current knowledge on mammalian parental care, focusing on its neural mechanisms, evolutionary origins, and derivatives. Neurobiological studies have identified specific neurons in the medial preoptic area as crucial for parental care. Unexpectedly, these neurons are characterized by the expression of molecules signaling satiety, such as calcitonin receptor and BRS3, and overlap with neurons involved in the reproductive behaviors of males but not females. A synthesis of comparative ecology and paleontology suggests an evolutionary scenario for mammalian parental care, possibly stemming from male-biased guarding of offspring in basal vertebrates. The terrestrial transition of tetrapods led to prolonged egg retention in females and the emergence of amniotes, skewing care toward females. The nocturnal adaptation of Mesozoic mammalian ancestors reinforced maternal care for lactation and thermal regulation via endothermy, potentially introducing metabolic gate control in parenting neurons. The established maternal care may have served as the precursor for paternal and cooperative care in mammals and also fostered the development of group living, which may have further contributed to the emergence of empathy and altruism. These evolution-informed working hypotheses require empirical validation, yet they offer promising avenues to investigate the neural underpinnings of mammalian social behaviors.

Shrub avoidance by an open-adapted ground squirrel in a shrub-encroached environment.

Habitat loss and degradation are leading drivers of the widespread decline in wildlife populations, and understanding how wildlife perceive and navigate their environment is useful for predicting responses to future landscape changes. Small mammals play an important role in their environments, however, many species are threatened by rapid environmental change. The Harris' antelope squirrel (Ammospermophilus harrisii) is endemic to the Sonoran Desert but faces multiple landscape changes due to anthropogenic activity. We fitted A. harrisii with radio collars to quantify resource selection and better understand how further environmental change may affect squirrels. Squirrels exhibited differential selection depending on behavior and scale. When selecting for microsites suitable for burrows and alarm calling (i.e., fourth-order selection), squirrels selected for both cacti and shrub portions of the habitat. Overall habitat selection within home ranges (i.e., third-order selection) showed selection against shrub patches, however, suggesting that further shrub encroachment may have consequences for A. harrisii behavior and distribution.

Seeking temporal refugia to heat stress: increasing nocturnal activity despite predation risk.

Flexibility in activity timing may enable organisms to quickly adapt to environmental changes. Under global warming, diurnally adapted endotherms may achieve a better energy balance by shifting their activity towards cooler nocturnal hours. However, this shift may expose animals to new or increased environmental challenges (e.g. increased predation risk, reduced foraging efficiency). We analysed a large dataset of activity data from 47 ibex (Capra ibex) in two protected areas, characterized by varying levels of predation risk (presence versus absence of the wolf-Canis lupus). We found that ibex increased nocturnal activity following warmer days and during brighter nights. Despite the considerable sexual dimorphism typical of this species and the consequent different predation-risk perception, males and females demonstrated consistent responses to heat in both predator-present and predator-absent areas. This supports the hypothesis that shifting activity towards nighttime may be a common strategy adopted by diurnal endotherms in response to global warming. As nowadays different pressures are pushing mammals towards nocturnality, our findings emphasize the urgent need to integrate knowledge of temporal behavioural modifications into management and conservation planning.

Leveraging limited data from wildlife monitoring in a conflict affected region in Venezuela.

Efficient monitoring of biodiversity-rich areas in conflict-affected areas with poor rule of law requires a combination of different analytical approaches to account for data biases and incompleteness. In the upland Amazon region of Venezuela, in Canaima National Park, we initiated biodiversity monitoring in 2015, but it was interrupted by the establishment of a large-scale mining development plan in 2016, compromising the temporal and geographical extent of monitoring and the security of researchers. We used a resource selection function model framework that considers imperfect detectability and supplemented detections from camera trap surveys with opportunistic off-camera records (including animal tracks and direct sighting) to (1) gain insight into the value of additional occurrence records to accurately predict wildlife resource use in the perturbated area (deforestation, fire, swidden agriculture, and human settlements vicinity), (2) when faced with security and budget constraints. Our approach maximized the use of available data and accounted for biases and data gaps. Adding data from poorly sampled areas had mixed results on estimates of resource use for restricted species, but improved predictions for widespread species. If budget or resources are limited, we recommend focusing on one location with both on-camera and off-camera records over two with cameras. Combining camera trap records with other field observations (28 mammals and 16 birds) allowed us to understand responses of 17 species to deforestation, 15 to fire, and 13 to swidden agriculture. Our study encourages the use of combinations of methods to support conservation in high-biodiversity sites, where access is restricted, researchers are vulnerable, and unequal sampling efforts exist.

Photoperiodic physiology of summer breeding birds and a search for the role of eye.

Photoperiod regulation of gonadal cycles is well studied and documented in both birds and mammals. Change in photoperiod is considered as the most effective and important cue to time the initiation of the annual physiological cycles in birds. Approaching of long days (as observed in summer months), signal long-day breeding birds to initiation reproduction and other related functions. Birds and other non-mammalian vertebrates use the extraocular photoreceptors which may be present in the mediobasal hypothalamus (MBH) or associated regions to measure the photoperiodic time and so are different from mammals where only the eyes are lone photoreceptive organs. The downstream signaling involves thyroid responsive genes playing a crucial role in mediating photoperiodic signals in both birds and mammals. Role of eyes in the avian seasonal cycle has been a questionable issue with evidences both favoring and negating any role. We propose that morphological as well as physiological data argue that retinal photoreceptors can participate in gonadal cycle, at least in the quail and duck. The present review details the studies of photoneuroendocrine control of gonadal axis in birds and review evidences to decipher the role eyes in photoperiodic mediated physiologies in birds.

Cardiorespiratory adaptations in small cetaceans and marine mammals.

The dive response, or the 'master switch of life', is probably the most studied physiological trait in marine mammals and is thought to conserve the available O2 for the heart and brain. Although generally thought to be an autonomic reflex, several studies indicate that the cardiovascular changes during diving are anticipatory and can be conditioned. The respiratory adaptations, where the aquatic breathing pattern resembles intermittent breathing in land mammals, with expiratory flow exceeding 160 litres s-1 has been measured in cetaceans, and where exposure to extreme pressures results in alveolar collapse (atelectasis) and recruitment upon ascent. Cardiorespiratory coupling, where breathing results in changes in heart rate, has been proposed to improve gas exchange. Cardiorespiratory coupling has also been reported in marine mammals, and in the bottlenose dolphin, where it alters both heart rate and stroke volume. When accounting for this respiratory dependence on cardiac function, several studies have reported an absence of a diving-related bradycardia except during dives that exceed the duration that is fuelled by aerobic metabolism. This review summarizes what is known about the respiratory physiology in marine mammals, with a special focus on cetaceans. The cardiorespiratory coupling is reviewed, and the selective gas exchange hypothesis is summarized, which provides a testable mechanism for how breath-hold diving vertebrates may actively prevent uptake of N2 during routine dives, and how stress results in failure of this mechanism, which results in diving-related gas emboli.

The longevity bottleneck hypothesis: Could dinosaurs have shaped ageing in present-day mammals?

The evolution and biodiversity of ageing have long fascinated scientists and the public alike. While mammals, including long-lived species such as humans, show a marked ageing process, some species of reptiles and amphibians exhibit very slow and even the absence of ageing phenotypes. How can reptiles and other vertebrates age slower than mammals? Herein, I propose that evolving during the rule of the dinosaurs left a lasting legacy in mammals. For over 100 million years when dinosaurs were the dominant predators, mammals were generally small, nocturnal, and short-lived. My hypothesis is that such a long evolutionary pressure on early mammals for rapid reproduction led to the loss or inactivation of genes and pathways associated with long life. I call this the 'longevity bottleneck hypothesis', which is further supported by the absence in mammals of regenerative traits. Although mammals, such as humans, can evolve long lifespans, they do so under constraints dating to the dinosaur era.

Functional consequences of animal community changes in managed grasslands: An application of the CAFE approach.

In the midst of an ongoing biodiversity crisis, much research has focused on species losses and their impacts on ecosystem functioning. The functional consequences (ecosystem response) of shifts in communities are shaped not only by changes in species richness, but also by compositional shifts that result from species losses and gains. Species differ in their contribution to ecosystem functioning, so species identity underlies the consequences of species losses and gains on ecosystem functions. Such research is critical to better predict the impact of disturbances on communities and ecosystems. We used the "Community Assembly and the Functioning of Ecosystems" (CAFE) approach, a modification of the Price equation to understand the functional consequences and relative effects of richness and composition changes in small nonvolant mammal and dung beetle communities as a result of two common disturbances in North American prairie restorations, prescribed fire and the reintroduction of large grazing mammals. Previous research in this system has shown dung beetles are critically important decomposers, while small mammals modulate much energy in prairie food webs. We found that dung beetle communities were more responsive to bison reintroduction and prescribed fires than small nonvolant mammals. Dung beetle richness increased after bison reintroduction, with higher dung beetle community biomass resulting from changes in remaining species (context-dependent component) rather than species turnover (richness components); prescribed fire caused a minor increase in dung beetle biomass for the same reason. For small mammals, bison reintroduction reduced energy transfer through the loss of species, while prescribed fire had little impact on either small mammal richness or energy transfer. The CAFE approach demonstrates how bison reintroduction controls small nonvolant mammal communities by increasing prairie food web complexity, and increases dung beetle populations with possible benefits for soil health through dung mineralization and soil bioturbation. Prescribed fires, however, have little effect on small mammals and dung beetles, suggesting a resilience to fire. These findings illustrate the key role of re-establishing historical disturbance regimes when restoring endangered prairie ecosystems and their ecological function.

Allometric scaling of skin weight and thickness to body weight in relation to taxonomic orders and habitats in mammals.

Skin is the largest organ in a mammal body, and it exhibits most significant range of adaptations to different habitats. It is a complex, biological composite structure, consisting of epidermis, dermis and subcutaneous tissues and is used for the therapeutic application of medical devices to improve healthcare. Extensive studies have been performed on the roles of the skin; however, little is known on its physiological characteristics in relation to body size among different species. The purpose of this study was therefore to evaluate the allometric scaling of skin weight (SW) and thickness (ST) to body weight (BW) in relation to genetics and habitats. Also analysed the relationship of BW to thicknesses of epidermis, dermis and subcutaneous tissues. This study used 249 adult animals of both sexes, belonging to 144 species, clustered in 18 taxonomic orders and five types of habitats. The animals were obtained from various sources in Japan. SW and BW were weighed, and ST was measured using a calliper followed by data analysis. Results showed that SW and ST were related to BW [log SW = 0.969 × logBW - 0636, adjust. R2 : 0.975]. The BW increased with increasing skin dermal thickness (y = 0.3916x + 1.5253, adjust. R2 : 0.6921), slightly with epidermal thickness (y = 0.2495x + 0.3984, adjust. R2 : 0.3402), but not all with the thickness of subcutaneous tissues (y = 0.1454x + 2.2437, adjust. R2 : 0.0752). The ratio of SW to BW (SW/BW) distributed over a large range from 0.06 to 0.64 values and varied among animal taxonomic orders and their dwelling habitats. Close relationship of BW to SW/BW was observed in species weighing ≥200 g but not in species weighing <200 g. In conclusion, SW and ST in mammals are determined by BW. The SW/BW varies based on BW, taxonomic orders and habitat and is large in small mammals weighing ≥200 g to provide a mechanism used for survival strategy.

A bird's eye view of the hippocampus beyond space: Behavioral, neuroanatomical, and neuroendocrine perspectives.

Although the hippocampus is one of the most-studied brain regions in mammals, research on the avian hippocampus has been more limited in scope. It is generally agreed that the hippocampus is an ancient feature of the amniote brain, and therefore homologous between the two lineages. Because birds and mammals are evolutionarily not very closely related, any shared anatomy is likely to be crucial for shared functions of their hippocampi. These functions, in turn, are likely to be essential if they have been conserved for over 300 million years. Therefore, research on the avian hippocampus can help us understand how this brain region evolved and how it has changed over evolutionary time. Further, there is a strong research foundation in birds on hippocampal-supported behaviors such as spatial navigation, food caching, and brood parasitism that scientists can build upon to better understand how hippocampal anatomy, network circuitry, endocrinology, and physiology can help control these behaviors. In this review, we summarize our current understanding of the avian hippocampus in spatial cognition as well as in regulating anxiety, approach-avoidance behavior, and stress responses. Although there are still some questions about the exact number of subdivisions in the avian hippocampus and how that might vary in different avian families, there is intriguing evidence that the avian hippocampus might have complementary functional profiles along the rostral-caudal axis similar to the dorsal-ventral axis of the rodent hippocampus, where the rostral/dorsal hippocampus is more involved in cognitive processes like spatial learning and the caudal/ventral hippocampus regulates emotional states, anxiety, and the stress response. Future research should focus on elucidating the cellular and molecular mechanisms - including endocrinological - in the avian hippocampus that underlie behaviors such as spatial navigation, spatial memory, and anxiety-related behaviors, and in so doing, resolve outstanding questions about avian hippocampal function and organization.

Melting climates shrink North American small mammals.

Mammals play important ecological roles in terrestrial ecosystems, with their particular niches and their impacts on energy flow and nutrient cycling being strongly influenced by one of their most fundamental traits-their body size. Body size influences nearly all of the physiological, behavioral, and ecological traits of mammals, and thus, shifts in body size often serve as key mechanisms of adaptation to variation in environmental conditions over space and time. Along with shifts in phenology and distributions, declining body size has been purported to be one of the three universal responses to anthropogenic climate change, yet few studies have been conducted at the spatial and temporal scales appropriate to test this claim. Here, we report that in response to warming of terrestrial ecosystems across North America over the past century, small mammals are decreasing in body size. We further estimate that by 2100 (when global temperatures may have risen some 2.5 to 5.5 °C since 1880), the total anthropogenic decline in body mass of these ecologically and economically important species may range from 10 to 21%. Such shifts in body size of the great multitudes of small mammal populations are, in turn, likely to have major impacts on the structural and functional diversity of terrestrial assemblages across the globe.

Endotherms trade body temperature regulation for the stress response.

Responding to perceived threats is energetically expensive and can require animals to curtail somatic repair, immunity, and even reproduction to balance energy ledgers. In birds and mammals, energetic demands of thermoregulation are often immense, yet whether homeostatic body temperatures are also compromised to aid the stress response is not known. Using data sourced from over 60 years of literature and 24 endotherm species, we show that exposure to non-thermal challenges (e.g. human interaction, social threats) caused body temperatures to decrease in the cold and increase in the warmth, but particularly when species-specific costs of thermoregulation were high and surplus energy low. Biophysical models revealed that allowing body temperature to change in this way liberated up to 24% (mean = 5%) of resting energy expenditure for use towards coping. While useful to avoid energetic overload, these responses nevertheless heighten risks of cold- or heat-induced damage, particularly when coincident with cold- or heatwaves.

Trait-based sensitivity of large mammals to a catastrophic tropical cyclone.

Extreme weather events perturb ecosystems and increasingly threaten biodiversity1. Ecologists emphasize the need to forecast and mitigate the impacts of these events, which requires knowledge of how risk is distributed among species and environments. However, the scale and unpredictability of extreme events complicate risk assessment1-4-especially for large animals (megafauna), which are ecologically important and disproportionately threatened but are wide-ranging and difficult to monitor5. Traits such as body size, dispersal ability and habitat affiliation are hypothesized to determine the vulnerability of animals to natural hazards1,6,7. Yet it has rarely been possible to test these hypotheses or, more generally, to link the short-term and long-term ecological effects of weather-related disturbance8,9. Here we show how large herbivores and carnivores in Mozambique responded to Intense Tropical Cyclone Idai, the deadliest storm on record in Africa, across scales ranging from individual decisions in the hours after landfall to changes in community composition nearly 2 years later. Animals responded behaviourally to rising floodwaters by moving upslope and shifting their diets. Body size and habitat association independently predicted population-level impacts: five of the smallest and most lowland-affiliated herbivore species declined by an average of 28% in the 20 months after landfall, while four of the largest and most upland-affiliated species increased by an average of 26%. We attribute the sensitivity of small-bodied species to their limited mobility and physiological constraints, which restricted their ability to avoid the flood and endure subsequent reductions in the quantity and quality of food. Our results identify general traits that govern animal responses to severe weather, which may help to inform wildlife conservation in a volatile climate.

Mechanical loading and hyperosmolarity as a daily resetting cue for skeletal circadian clocks.

Daily rhythms in mammalian behaviour and physiology are generated by a multi-oscillator circadian system entrained through environmental cues (e.g. light and feeding). The presence of tissue niche-dependent physiological time cues has been proposed, allowing tissues the ability of circadian phase adjustment based on local signals. However, to date, such stimuli have remained elusive. Here we show that daily patterns of mechanical loading and associated osmotic challenge within physiological ranges reset circadian clock phase and amplitude in cartilage and intervertebral disc tissues in vivo and in tissue explant cultures. Hyperosmolarity (but not hypo-osmolarity) resets clocks in young and ageing skeletal tissues and induce genome-wide expression of rhythmic genes in cells. Mechanistically, RNAseq and biochemical analysis revealed the PLD2-mTORC2-AKT-GSK3ß axis as a convergent pathway for both in vivo loading and hyperosmolarity-induced clock changes. These results reveal diurnal patterns of mechanical loading and consequent daily oscillations in osmolarity as a bona fide tissue niche-specific time cue to maintain skeletal circadian rhythms in sync.

Collagen IV of basement membranes: II. Emergence of collagen IVα345 enabled the assembly of a compact GBM as an ultrafilter in mammalian kidneys.

The collagen IVα345 (Col-IVα345) scaffold, the major constituent of the glomerular basement membrane (GBM), is a critical component of the kidney glomerular filtration barrier. In Alport syndrome, affecting millions of people worldwide, over two thousand genetic variants occur in the COL4A3, COL4A4, and COL4A5 genes that encode the Col-IVα345 scaffold. Variants cause loss of scaffold, a suprastructure that tethers macromolecules, from the GBM or assembly of a defective scaffold, causing hematuria in nearly all cases, proteinuria, and often progressive kidney failure. How these variants cause proteinuria remains an enigma. In a companion paper, we found that the evolutionary emergence of the COL4A3, COL4A4, COL4A5, and COL4A6 genes coincided with kidney emergence in hagfish and shark and that the COL4A3 and COL4A4 were lost in amphibians. These findings opened an experimental window to gain insights into functionality of the Col-IVα345 scaffold. Here, using tissue staining, biochemical analysis and TEM, we characterized the scaffold chain arrangements and the morphology of the GBM of hagfish, shark, frog, and salamander. We found that α4 and α5 chains in shark GBM and α1 and α5 chains in amphibian GBM are spatially separated. Scaffolds are distinct from one another and from the mammalian Col-IVα345 scaffold, and the GBM morphologies are distinct. Our findings revealed that the evolutionary emergence of the Col-IVα345 scaffold enabled the genesis of a compact GBM that functions as an ultrafilter. Findings shed light on the conundrum, defined decades ago, whether the GBM or slit diaphragm is the primary filter.

Climate change should drive mammal defaunation in tropical dry forests.

Human-induced climate change has intensified negative impacts on socioeconomic factors, the environment, and biodiversity, including changes in rainfall patterns and an increase in global average temperatures. Drylands are particularly at risk, with projections suggesting they will become hotter, drier, and less suitable for a significant portion of their species, potentially leading to mammal defaunation. We use ecological niche modelling and community ecology biodiversity metrics to examine potential geographical range shifts of non-volant mammal species in the largest Neotropical dryland, the Caatinga, and evaluate impacts of climate change on mammal assemblages. According to projections, 85% of the mammal species will lose suitable habitats, with one quarter of species projected to completely lose suitable habitats by 2060. This will result in a decrease in species richness for more than 90% of assemblages and an increase in compositional similarity to nearby assemblages (i.e., reduction in spatial beta diversity) for 70% of the assemblages. Small-sized mammals will be the most impacted and lose most of their suitable habitats, especially in highlands. The scenario is even worse in the eastern half of Caatinga where habitat destruction already prevails, compounding the threats faced by species there. While species-specific responses can vary with respect to dispersal, behavior, and energy requirements, our findings indicate that climate change can drive mammal assemblages to biotic homogenization and species loss, with drastic changes in assemblage trophic structure. For successful long-term socioenvironmental policy and conservation planning, it is critical that findings from biodiversity forecasts are considered.

Field physiology in the aquatic realm: ecological energetics and diving behavior provide context for elucidating patterns and deviations.

Comparative physiology has developed a rich understanding of the physiological adaptations of organisms, from microbes to megafauna. Despite extreme differences in size and a diversity of habitats, general patterns are observed in their physiological adaptations. Yet, many organisms deviate from the general patterns, providing an opportunity to understand the importance of ecology in determining the evolution of unusual adaptations. Aquatic air-breathing vertebrates provide unique study systems in which the interplay between ecology, physiology and behavior is most evident. They must perform breath-hold dives to obtain food underwater, which imposes a physiological constraint on their foraging time as they must resurface to breathe. This separation of two critical resources has led researchers to investigate these organisms' physiological adaptations and trade-offs. Addressing such questions on large marine animals is best done in the field, given the difficulty of replicating the environment of these animals in the lab. This Review examines the long history of research on diving physiology and behavior. We show how innovative technology and the careful selection of research animals have provided a holistic understanding of diving mammals' physiology, behavior and ecology. We explore the role of the aerobic diving limit, body size, oxygen stores, prey distribution and metabolism. We then identify gaps in our knowledge and suggest areas for future research, pointing out how this research will help conserve these unique animals.

Energetic costs of feeding in 12 species of small-bodied primates.

There are no comparative, empirical studies of the energetic costs of feeding in mammals. As a result, we lack physiological data to better understand the selection pressures on the mammalian feeding apparatus and the influence of variables such as food geometric and material properties. This study investigates interspecific scaling of the net energetic costs of feeding in relation to body size, jaw-adductor muscle mass and food properties in a sample of 12 non-human primate species ranging in size from 0.08 to 4.2 kg. Net energetic costs during feeding were measured by indirect calorimetry for a variety of pre-cut and whole raw foods varying in geometric and material properties. Net feeding costs were determined in two ways: by subtracting either the initial metabolic rate prior to feeding or subtracting the postprandial metabolic rate. Interspecific scaling relationships were evaluated using pGLS and OLS regression. Net feeding costs scale negatively relative to both body mass and jaw-adductor mass. Large animals incur relatively lower feeding costs indicating that small and large animals experience and solve mechanical challenges in relation to energetics in different ways. This article is part of the theme issue 'Food processing and nutritional assimilation in animals'.