Short-term dynamics of beaver dam flow states.

Beavers (Castor canadensis and C. fiber) build dams that modify catchment and pond water balances, and it has been suggested that they can be a nature-based solution for reducing flood hydrographs, enhancing low flow hydrographs and restoring hydrological functioning of degraded streams. How water moves through a beaver dam is determined by its flow state (e.g., overflow, underflow). However, current conceptual models only consider flow state as changing over the beaver site occupation-abandonment cycle. To assess whether flow state changes at shorter timescales and identify possible triggers (e.g., rainfall, animals), we integrated camera trap imagery, machine learning, water level measurements, and hydrometeorological data at beaver dams in a montane peatland in the Canadian Rocky Mountains. Contrary to current models, we found that flow states changed frequently, changing a maximum 12 times during the 139-day study period, but that changes had limited synchronicity amongst the dams in the same stream. More than two-thirds of the changes coincided with rainfall events. We observed no changes in flow state in response to beaver activity or wildlife crossings perhaps due to the camera positioning. Our findings augment the long-term oriented framework, which links changes to the occupancy cycle of a beaver pond and frequent and hydrological-driven changes. To develop realistic predictions of hydrological impacts of beaver dams, ecohydrological models should update their representation of the influence of beaver dams to include short-term dynamism of flow states and potential triggers. Our study advances the understanding of the important, yet understudied, role of beaver dams in stream restoration and climate change initiatives.

Effects of acute maternal stress induced by predator cues on spatial learning and memory of offspring in the subterranean rodent Ctenomys talarum.

One of the main selection pressures to which animals are exposed in nature is predation, which affects a wide variety of biological traits. When the mother experiences this stressor during pregnancy and/or lactation, behavioral and physiological responses may be triggered in the offspring as well. Thus, in order to broaden and deepen knowledge on the transgenerational effects of predation stress, we evaluated how maternal stress experienced during pregnancy and/or lactation affects the spatial abilities of progeny at the onset of adulthood in the subterranean rodent Ctenomys talarum. The results showed that, contrary to what was observed in other rodent species, maternal exposure to predator cues during pregnancy and lactation did not negatively affect the spatial abilities of the offspring, even registering some minor positive effects. Concomitantly, no effects of predatory cues on physiological parameters associated with stress were observed in the progeny. This difference in results between the present study and previous works on maternal stress highlights the importance of considering the species to be evaluated (strain, age and origin-wild or captive-) and the type of stressor used (artificial or natural, intensity of exposure) in the evaluation of the possible transgenerational effects of maternal stress.

Embryonic-placental relationship in Lagostomus maximus as compared to other hystricognath rodents and eutherian mammals.

Reproductive specializations in caviomorphs (infraorder Hystricognathi), are very peculiar within the Order Rodentia. These include long gestations, the birth of offspring with an extreme degree of precociality, and short lactation periods. This study describes the embryo-placental relationship of viable implantation sites (IS) of the plains viscacha, Lagostomus maximus, after 46 post-coital days. The observations recorded in this study are comparatively discussed with those of other hystricognaths and eutherians. At this stage, the embryo resembles that of other eutherians. At this time of embryo development, the placenta exhibits a size, shape, and organization similar to that it will have in its mature state. Besides, the subplacenta is already highly folded. These characteristics are adequate to sustain the development of future precocial offspring. The mesoplacenta, a structure present in other hystricognaths and related to uterine regeneration is described for the first time in this species. This detailed description of the placental and embryonic structure contributes to the knowledge of the reproductive and developmental biology of the viscacha, as well as that of hystricognaths. These characteristics will allow testing other hypotheses related to the morphology and physiology of the placenta and subplacenta, and their relationship with the growth and development of precocial offspring in Hystricognathi.

A nonparametric test of group distributional differences for hierarchically clustered functional data.

Biological sex and gender are critical variables in biomedical research, but are complicated by the presence of sex-specific natural hormone cycles, such as the estrous cycle in female rodents, typically divided into phases. A common feature of these cycles are fluctuating hormone levels that induce sex differences in many behaviors controlled by the electrophysiology of neurons, such as neuronal membrane potential in response to electrical stimulus, typically summarized using a priori defined metrics. In this paper, we propose a method to test for differences in the electrophysiological properties across estrous cycle phase without first defining a metric of interest. We do this by modeling membrane potential data in the frequency domain as realizations of a bivariate process, also depending on the electrical stimulus, by adopting existing methods for longitudinal functional data. We are then able to extract the main features of the bivariate signals through a set of basis function coefficients. We use these coefficients for testing, adapting methods for multivariate data to account for an induced hierarchical structure that is a product of the experimental design. We illustrate the performance of the proposed approach in simulations and then apply the method to experimental data.

Effects of water restriction and dirt on grooming behavior in neotropical rodents (Trinomys setosus and T. yonenagae) (Echimyidae).

Grooming in rodents presents an evolutionarily conserved behavioral pattern that may cause water loss since saliva is deposited during washing onto large body surfaces. Trinomys yonenagae and Trinomys setosus are sister species of spiny rats occurring in Brazil, the former inhabiting a paleodesert of fixed dunes in the Caatinga, the latter being found in mesic environments of the Atlantic Forest. Consequently, it is expected that both species evolved under different selective pressures related to water balance, with T. yonenagae presenting mechanisms for dealing with water deprivation not found in T. setosus. Reduction of self-cleaning expression seems to offer a possible way to save water, as previously suggested by studies of the sand-dwelling spiny rat. Therefore, we propose to investigate grooming under four conditions: 'control' (C), a regimen of 'water restriction' (WR), of 'dirt' (D), and the combination of both conflicting stimuli (WR + D), in T. setosus, T. yonenagae, and Rattus norvegicus to compare the behavioral responses of these species. The main differences are observed in the forest dweller: T. setosus expresses a low relative duration of face washing under C, whose value is intermediate between the ones found in the two other species. WR treatment does not alter this pattern, however, the addition of dirt (D, WR + D) significantly increases the relative duration of washing in relation to C. Locomotor activity is decreased both in T. setosus and Wistar rats when they are under WR, a situation that could jeopardize antipredatory performance. T. yonenagae, the sand dweller, maintains a significantly lower expression of washing under C, as previously suggested, and under WR, D and WR + D. In addition, differently from the other two species the sand dweller maintains a normal activity level during all treatments. This study suggests differences in grooming as a strategy alluding to water balance by the two spiny rats inhabiting different ecosystems. A significantly clear pattern that saves water is observed in T. yonenagae, which probably has contributed to his evolution in one of the hottest semiarid areas of the world.

Rodent responses to volatile compounds provide insights into the function of floral scent in mammal-pollinated plants.

Flowers pollinated by mammals have evolved in many plant families. Several scent compounds that attract bats to flowers have been identified, but the chemical ecology of pollination mutualisms between plants and ground-dwelling mammals is poorly understood. Rodents are key pollinators in South Africa and rely heavily on olfaction to locate food. Our aim was to identify compounds that may function to attract rodents to flowers. Eighteen volatile compounds, including 14 that are prominent in the scent of rodent-pollinated flowers, were used in choice experiments involving wild-caught individuals of four native rodent species. Rodents were generally attracted to oxygenated aliphatic compounds, specifically ketones and esters, but not to some aromatic compounds common in floral scents of insect-pollinated species, nor to a sulfide compound that is attractive to bats. Associative conditioning using sugar solution as a reward had only weak effects on the attractiveness of compounds to rodents. The attractive effect of some compounds disappeared when they were blended with compounds that did not attract rodents. We conclude that aliphatic ketones and esters are likely to play a key role in attracting rodents to flowers. Deployment of these compounds may allow plants to exploit rodent sensory bias that evolved in other contexts such as intra-specific communication and searching for seeds. This article is part of the theme issue 'Natural processes influencing pollinator health: from chemistry to landscapes'.

How beavers affect riverine aquatic macroinvertebrates: a review.

Background: As ecosystem engineers, the construction of dams by beavers alters stream habitat physically and biologically, making them a species of interest for habitat restoration. Beaver-created habitat changes affect a wide range of aquatic invertebrate species. However, despite numerous individual studies of how beavers affect aquatic macroinvertebrate assemblages, there has been no evaluation of the consensus of these effects across studies. Methodology: We collated and examined studies comparing beaver-created ponds to nearby lotic reaches to determine general trends in aquatic macroinvertebrate richness, density, biomass, and functional composition between habitats. From this evidence, we highight knowledge gaps in how beaver activity affects aquatic macroinvertebrates. Results: Overall, in the majority of studies, aquatic macroinvertebrate richness was higher in nearby lotic reaches compared to beaver-created ponds, but richness at coarser scales (gamma diversity) increased with the addition of beaver ponds due to increased habitat heterogeneity. Functional feeding group (FFG) patterns were highly context-dependent, though predator taxa were generally more abundant in beaver ponds than adjacent lotic reaches. Site-specific geomorphological changes, coupled with dam or riparian zone characteristics and resulting differences in basal food resources likely shape other FFG responses. Conclusions: We identify a lack of long-term studies at single or multiple sites and conclude that fine-scale approaches may improve our understanding of the dynamics of macroinvertebrates within the freshwater realm and beyond. Due to the context-dependent nature of each study, further systematic studies of beaver engineering effects across a wider variety of environmental conditions and wetland types will also help inform land and species management decisions, such as where to prioritize protection of beaver habitats in the face of a global freshwater biodiversity crisis, or where to restore beaver populations to deliver maximum benefit.

Sex, not social behavior, predicts fecal glucocorticoid metabolite concentrations in a facultatively social rodent, the highland tuco-tuco (Ctenomys opimus).

Social relationships may influence circulating glucocorticoid levels, particularly in group-living species in which individuals regularly engage in interactions with conspecifics. The effects of such interactions appear to vary, with greater social contact being associated with increased glucocorticoid concentrations in some species but decreased concentrations in others. These distinct responses raise intriguing questions regarding relationships among social behavior, individual phenotypes, and glucocorticoid physiology. To explore such relationships in a free-living mammal with a dynamic social organization, we quantified variation in baseline glucocorticoids in a population of highland tuco-tucos (Ctenomys opimus) from Jujuy Province, Argentina. These subterranean rodents are facultatively social, with lone and group-living individuals regularly occurring within the same population. To assess potential endocrine correlates of this behavioral variability, we examined differences in baseline fecal glucocorticoid metabolite (fGCm) concentrations as a function of social group size and composition as well as several metrics of social behavior derived from social network analyses. Despite marked variability in social relationships among the 37 (12 male, 25 female) free-living tuco-tucos sampled, none of the measures of social behavior examined were significant predictors of variation in fGCm concentrations. In contrast, individual variation in glucocorticoid metabolites was best explained by sex, with males having higher fGCm concentrations than females. These analyses provide the first characterization of the glucocorticoid physiology of highland tuco-tucos and underscore the potential importance of intrinsic phenotypic factors (e.g., sex) in shaping glucocorticoid variation in free-living mammals.

Facilitative interaction promotes occupancy of a desert amphibian across a climate gradient.

Biotic interactions and environmental factors influence a species' occurrence. Facilitative interactions have the potential to expand species occupancy and mitigate abiotic stress, but are often not considered. The Great Basin clade of the Columbia spotted frog (Rana luteiventris) is an aquatic amphibian found in the arid and semi-arid Great Basin, USA. Aquatic resources in this system are limited and likely to change under future climate scenarios. American beavers (Castor canadensis) are ecosystem engineers that impact the surrounding landscape by increasing water availability. Consequently, beaver-created habitat can facilitate the presence of other species by providing the aquatic resources required by many aquatic and riparian animals. Our objective was to understand patterns of co-occurrence between Columbia spotted frogs and beavers across environmental gradients in the Great Basin, USA. We used environmental DNA detections to quantify the co-occurrence of Columbia spotted frog and beaver using a two-species occupancy framework. At the lowest annual precipitation (202 mm), Columbia spotted frogs were 3× (95% CI 2.35, 3.96) more likely to occur with beavers than without; however, they were less likely to occur with beavers once precipitation was > 380 mm. Thus, beaver activity may increase aquatic resources for Columbia spotted frogs in the Great Basin that may mitigate changing precipitation patterns under climate change. Facilitative interactions likely extend to other aquatic species in arid systems and highlight an important consideration for ecologists when evaluating a species' response to climate change, and may promote the formation of refugia for species with strict abiotic tolerances and dispersal limitations.

Autonomic control of pancreatic beta cells: What is known on the regulation of insulin secretion and beta-cell proliferation in rodents and humans.

Insulin secretion and pancreatic beta-cell proliferation are tightly regulated by several signals such as hormones, nutrients, and neurotransmitters. However, the autonomic control of beta cells is not fully understood. In this review, we describe mechanisms involved in insulin secretion as well as metabolic and mitogenic actions on its target tissues. Since pancreatic islets are physically connected to the brain by nerves, parasympathetic and sympathetic neurotransmitters can directly potentiate or repress insulin secretion and beta-cell proliferation. Finally, we highlight the role of the autonomic nervous system in metabolic diseases such as diabetes and obesity.

The impact of bio-logging on body weight change of the Eurasian beaver.

Bio-logging is a common method to collect ecological data on wild animals, but might also induce stress, reduce body condition, and alter behavior. Eurasian beavers (Castor fiber) are a semi-aquatic and nocturnal species that are challenging to observe in the wild. Bio-loggers are hence useful tools to study their behaviour and movements, but this raises concerns of potential negative impacts of tagging. To investigate the potential negative impacts of glue-on tags, we compared body weight change for tagged and untagged Eurasian beavers. We hypothesized that tagged beavers would gain less body weight compared to untagged beavers, and that weight change might be affected by tagging length, tag weight, water temperature and the season of tagging. Daily percentage body weight change in relation to initial body weight during the first capture was compared during 57 tagging periods (18±7 days) and 32 controls periods (64±47 days). Body weight change varied between the two groups, with untagged beavers on average gaining daily weight whilst tagged beavers on average lost weight daily, indicating a negative effect of tagging. The average reduction in percentage body weight change per day for tagged beavers was small (0.1 ± 0.3%), and with large individual variation. Neither tag weight, number of tagging days, nor season were important in explaining body weight change of tagged animals. In other words, we found that tagging reduced daily body weight during the tagging period but were unable to determine the mechanism(s) responsible for this decline. Detrimental effects of tagging have important implications for animal welfare and can introduce bias in data that are collected. This calls for careful consideration in the use of tags. We conclude that studies investigating the effects of tagging should consider individual variation in the effects of tagging and, where possible, compare tagged animals with a control group.

Seed traits and rodent community interact to determine seed fate: evidence from both enclosure and field experiments.

Animal-mediated seed dispersal is an important ecological process in which a strong mutualism between animals and plants can arise. However, few studies have examined how a community of potential seed dispersers interacts with sympatric seed trees. We employed a series of experiments in the Qinling Mountains in both semi-natural enclosure and the field to assess the interactions among 3 sympatric rodent species and 3 Fagaceae tree seeds. Seed traits all showed similar tannin levels but markedly different physical traits and nutritional contents. We found that seeds with heavy weight, thick coat, and high nutritional contents were less likely to be eaten in situ but more often to be eaten after dispersal or hoarded by rodents. These results support both the handling time hypothesis and the high nutrition hypothesis. Surprisingly, we also found that rodents, maybe, preferred to consume seeds with low levels of crude fiber in situ, and to harvest and hoard those with high levels of crude fiber for later consumption. The sympatric rodent species, Cansumys canus, the largest rodent in our study, harvested and hoarded more Quercus variabilis seeds with high physical and nutritional traits, while Apodemus draco, the smallest rodent, harvested more Q. serrata seeds with low physical and nutritional traits, and Niviventer confucianus harvested and hoarded more Q. aliena seeds with medium physical and nutritional traits. Our study demonstrates that different seed traits play different roles in influencing the seed fate and the shaping of mutualism and predation interactions within a community of rodent species.

Increased burrow oxygen levels trigger defensive burrow-sealing behavior by plateau zokors.

Defensive behaviors are a response to immediate and potential threats in the environment, including abiotic and biotic threats. Subterranean rodents exhibit morphological and physiological adaptions for life underground, and they will seal with mounds and additional plugs when their burrow opened. However, little is known about the factors driving this defensive behavior. In this study, we selected a subterranean rodent, plateau zokor (Myospalax fontanieri), as a species to investigate (both in the laboratory and in the field) the possible factors responsible for burrow-sealing behavior. Our results showed that: (1) In the laboratory, the burrow-sealing frequency of plateau zokor in response to five factors were as follows: oxygen (52.63%) > light (34.58%) > temperature (20.24%) > gas flow (6.48%) > sound/control (0%). Except for light, the burrow-sealing frequency in response to other factors was significantly lower than that in response to oxygen (P < 0.05). (2) Burrow-sealing behavior in response to each treatment did not differ significantly between males and females in the laboratory experiment. (3) In the field, during the animal's active periods in both the cold and warm season, the burrow-sealing frequency under the oxygen treatment was higher than that under the light and temperature treatments. Plateau zokors were found not to be sensitive to these treatments during their inactive periods during both the cold and warm season. (4) The latency to reseal the burrow showed no obvious differences between each treatment both in the laboratory and in the field. In conclusion, the main factor that influences the burrow-sealing behavior of plateau zokors is the variation in oxygen concentration, and this defensive behavior is related to their activity rhythm.

Phylogenomic relationships and molecular convergences to subterranean life in rodent family Spalacidae.

All extant species in the rodent family Spalacidae are subterranean and have evolved various traits for underground life. However, the phylogenomic relationships among its three subfamilies (Myospalacinae, Spalacinae, and Rhizomyinae) and the molecular basis underlying their adaptations to underground life remain poorly understood. Here, we inferred the phylogenomic relationships among these subfamilies based on de novo sequencing the genome of the hoary bamboo rat ( Rhizomys pruinosus). Analyses showed that ~50% of the identified 11 028 one-to-one orthologous protein-coding genes and the concatenated sequences of these orthologous genes strongly supported a sister relationship between Myospalacinae and Rhizomyinae. The three subfamilies diversified from each other within ~2 million years. Compared with the non-subterranean controls with similar divergence dates, the spalacids shared more convergent genes with the African subterranean mole-rats at the genomic scale due to more rapid protein sequence evolution. Furthermore, these convergent genes were enriched in the functional categories of carboxylic acid transport, vascular morphogenesis, and response to oxidative stress, which are closely associated with adaptations to the hypoxic-hypercapnic underground environment. Our study presents a well-supported phylogenomic relationship among the three subfamilies of Spalacidae and offers new insights into the molecular adaptations of spalacids living underground.

Assessment of crop damage by rodent pests from experimental barley crop fields in Farta District, South Gondar, Ethiopia.

This study was conducted in Farta district, south Gondar from 2019 to 2020 cropping years to identify rodent pest species and estimate damage caused on barley crops. Four independent barley crop fields (40 x 40 m each) were sampled randomly to estimate the loss. Two were located near Alemsaga Priority State Forest and the other two were away from the forest. Four (2 x 2 m) rodent exclusion plots were established at 10 m interval as control units in each selected experimental barley fields using fine wire mesh. Rodent pest species were collected using both Sherman and snap traps throughout the different crop growing stages. The damaged and undamaged barley tillers by pest rodents were counted on five 1 x 1 m randomly sampled quadrats for each selected experimental fields. Variations on pest rodent population between cropping years and sites were analyzed using Chi square test. The mean crop damages between cropping years and experimental field sites were analyzed using two way ANOVA. Arvicanthis abyssinicus, Mastomys natalensis, Arvicanthis dembeensis, Mus musculus, Lophuromys simensis, Tachyoryctes splendens and Hystrix cristata were identified as pest rodents in the study area. A total of 968 individual rodents (427 in 2019 and 541 in 2020) were trapped during the study period. There was a statistical variation (χ2 = 13.42, df = 1 and P<0.05) between trapped individuals of the two successive years. The crop fields near the forest were more vulnerable than away from the forest during both cropping years. Statistical variations was observed on mean crop losses between cropping years and experimental barley crop sites. The highest crop damage was seen at maturity stage and the lowest during sowing in all experimental plots and cropping years. The percentage of barley yield loss due to rodent pests was 21.7 kg ha-1. The monetary value of this yield loss was equivalent to 4875 Birr (121.9 US$ h-1). Alemsaga Forest as shelter and conservation strategies like free of farmland from livestock and terracing for soil conservation have great role for the high rodent pest populations in the study area. Field sanitation, trapping and using restricted rodenticides like zinc phosphide are the possible recommendation to local farmers against rodent pests.

The effect of feed restriction, sex and age on the carcass composition and meat quality of nutrias (Myocastor coypus).

The objective of the study was to evaluate the effect of quantitative feed restriction on the carcass composition and physical, chemical, histochemical and sensory properties of meat from male and female nutria at the end of the feed restriction and fattening period. In the experiment, from two to eight months of age, males and females were divided into two groups: one group was fed ad libitum throughout the experiment, and the second group was restricted to two weeks (from 11 to 12 weeks of age). The restricted nutrias received 75% ad libitum feeding. The significant interaction among feed restriction, sex and age was revealed only in the cross-sectional area of all muscle fiber types. Age was the main factor affecting carcass composition and meat quality characteristics. Sex-related differences in carcass composition and some sensory attributes and feed restriction-related differences in carcass composition were observed.

Rare crested rat subfossils unveil Afro-Eurasian ecological corridors synchronous with early human dispersals.

Biotic interactions between Africa and Eurasia across the Levant have invoked particular attention among scientists aiming to unravel early human dispersals. However, it remains unclear whether behavioral capacities enabled early modern humans to surpass the Saharo-Arabian deserts or if climatic changes triggered punctuated dispersals out of Africa. Here, we report an unusual subfossil assemblage discovered in a Judean Desert's cliff cave near the Dead Sea and dated to between ∼42,000 and at least 103,000 y ago. Paleogenomic and morphological comparisons indicate that the specimens belong to an extinct subspecies of the eastern African crested rat, Lophiomys imhausi maremortum subspecies nova, which diverged from the modern eastern African populations in the late Middle Pleistocene ∼226,000 to 165,000 y ago. The reported paleomitogenome is the oldest so far in the Levant, opening the door for future paleoDNA analyses in the region. Species distribution modeling points to the presence of continuous habitat corridors connecting eastern Africa with the Levant during the Last Interglacial ∼129,000 to 116,000 y ago, providing further evidence of the northern ingression of African biomes into Eurasia and reinforcing previous suggestions of the critical role of climate change in Late Pleistocene intercontinental biogeography. Furthermore, our study complements other paleoenvironmental proxies with local-instead of interregional-paleoenvironmental data, opening an unprecedented window into the Dead Sea rift paleolandscape.

Echolocation in soft-furred tree mice.

Echolocation is the use of reflected sound to sense features of the environment. Here, we show that soft-furred tree mice (Typhlomys) echolocate based on multiple independent lines of evidence. Behavioral experiments show that these mice can locate and avoid obstacles in darkness using hearing and ultrasonic pulses. The proximal portion of their stylohyal bone fuses with the tympanic bone, a form previously only seen in laryngeally echolocating bats. Further, we found convergence of hearing-related genes across the genome and of the echolocation-related gene prestin between soft-furred tree mice and echolocating mammals. Together, our findings suggest that soft-furred tree mice are capable of echolocation, and thus are a new lineage of echolocating mammals.

Responses of small mammals to habitat characteristics in Southern Carpathian forests.

Compared to Northern Carpathians, the small mammal fauna of Southern Carpathian forests is poorly known, with no data on habitat use; our study seeks to fill this gap. To this end, we conducted a survey in the Southern Carpathians for five years, assessing habitat use by small mammals in forests along an elevational gradient. Trapping was done using live traps set in transects at elevations between 820 and 2040 m. For each transect we evaluated variables related to vegetation structure, habitat complexity, and geographical location. We considered abundance, species composition and species richness as response variables. The rodents Apodemus flavicollis and Myodes glareolus and the shrew Sorex araneus were common and dominant. Their abundance were positively correlated with tree cover, the best explanatory variable. Responses to other variables were mixed. The strong divergence in the relative habitat use by the three most abundant species may act as a mechanism that enables their coexistence as dominant species, exploiting the same wide range of habitat resources. Overall, habitat use in our study area was similar to that reported from Northern Carpathians, but we found also important differences probably caused by the differences in latitude and forest management practices.

The CPGs for Limbed Locomotion-Facts and Fiction.

The neuronal networks that generate locomotion are well understood in swimming animals such as the lamprey, zebrafish and tadpole. The networks controlling locomotion in tetrapods remain, however, still enigmatic with an intricate motor pattern required for the control of the entire limb during the support, lift off, and flexion phase, and most demandingly when the limb makes contact with ground again. It is clear that the inhibition that occurs between bursts in each step cycle is produced by V2b and V1 interneurons, and that a deletion of these interneurons leads to synchronous flexor-extensor bursting. The ability to generate rhythmic bursting is distributed over all segments comprising part of the central pattern generator network (CPG). It is unclear how the rhythmic bursting is generated; however, Shox2, V2a and HB9 interneurons do contribute. To deduce a possible organization of the locomotor CPG, simulations have been elaborated. The motor pattern has been simulated in considerable detail with a network composed of unit burst generators; one for each group of close synergistic muscle groups at each joint. This unit burst generator model can reproduce the complex burst pattern with a constant flexion phase and a shortened extensor phase as the speed increases. Moreover, the unit burst generator model is versatile and can generate both forward and backward locomotion.