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194 results for “Competitive effect”
Data from: Effect of plant root symbionts on performance of native woody species in competition with an invasive grass in multispecies microcosms
The majority of terrestrial plants form mutualistic associations with arbuscular mycorrhizal fungi (AMF) and rhizobia (i.e. nitrogen fixing bacteria). Understanding these associations has important implications for ecological theory and for restoration practice. Here we tested whether the presence of AMF and rhizobia influence the performance of native woody plants invaded by a non-native grass in experimental microcosms. We planted eight plant species (i.e. Acacia acuminata, A. microbotrya, Eucalyptus loxophleba subsp. loxophleba, E. astringens, Calothamnus quadrifidus, Callistemon phoeniceus, Hakea lissocarpha and H. prostrata) in microcosms of field-conditioned soil with and without addition of AMF and rhizobia in a fully factorial experimental design. After seedling establishment, we seeded half the microcosms with an invasive grass Bromus diandrus. We measured shoot and root biomass of native plants and Bromus, and on roots, the percentage colonization by AMF, number of rhizobia-forming nodules and number of proteaceous root clusters. We found no effect of plant root symbionts or Bromus addition on performance of myrtaceous, and as predicted, proteaceous species as they rely little or not at all on AMF and rhiozbia. Soil treatments with AMF and rhiozbia had a strong positive effect (i.e. larger biomass) on native legumes (A. microbotrya and A. acuminata). However, the beneficial effect of root symbionts on legumes became negative (i.e. lower biomass and less nodules) if Bromus was present, especially for one legume, i.e. A. acuminata, suggesting a disruptive effect of the invader on the mutualism. We also found a stimulating effect of Bromus on root nodule production in A. microbotrya and AMF colonization in A. acuminata which could be indicative of legumes' increased resource acquisition requirement, i.e. for nitrogen and phosphorus, respectively, in response to the Bromus addition. We have demonstrated the importance of measuring belowground effects because the aboveground effects gave limited indication of the effects occuring belowground.
Data from: Lions and leopards coexist without spatial, temporal or demographic effects of interspecific competition
1. Although interspecific competition plays a principle role in shaping species behaviour and demography, little is known about the population-level outcomes of competition between large carnivores, and the mechanisms that facilitate coexistence. 2. We conducted a multi-landscape analysis of two widely distributed, threatened large carnivore competitors to offer insight into coexistence strategies and assist with species-level conservation. 3. We evaluated how interference competition affects occupancy, temporal activity and population density of a dominant competitor, the lion (Panthera leo), and its subordinate competitor, the leopard (Panthera pardus). We collected camera-trap data over three years in ten study sites covering 5,070 km2. We used multispecies occupancy modelling to assess spatial responses in varying environmental and prey conditions and competitor presence, and examined temporal overlap and the relationship between lion and leopard densities across sites and years. 4. Results showed that both lion and leopard occupancy was independent of – rather than conditional on – their competitor's presence across all environmental covariates. Marginal occupancy probability for leopard was higher in areas with more bushy, 'hideable' habitat, human (tourist) activity and topographic ruggedness, whereas lion occupancy decreased with increasing hideable habitat and increased with higher abundance of very large prey. Temporal overlap was high between carnivores and there was no detectable relationship between species densities. 4. Lions pose a threat to the survival of individual leopards, but they exerted no tractable influence on leopard spatial or temporal dynamics. Furthermore, lions did not appear to suppress leopard populations, suggesting that intraguild competitors can coexist in the same areas without population decline. Aligned conservation strategies that promote functioning ecosystems, rather than target individual species, are therefore advised to achieve cost- and space-effective conservation.
Data from: The effect of elevated CO2 on growth and competition in experimental phytoplankton communities
We report an experiment designed to identify the effect of elevated CO2 on species of phytoplankton in a simple laboratory system. Major taxa of phytoplankton differ in their ability to take up CO2, which might lead to predictable changes in the growth rate of species and thereby shifts in the composition of phytoplankton communities in response to rising CO2. Six species of phytoplankton belonging to three major taxa (cyanobacteria, diatoms and chlorophytes) were cultured in atmospheres whose CO2 concentration was gradually increased from ambient levels to 1000 parts per million over about 100 generations and then maintained for a further 200 generations at elevated CO2. The experimental design allowed us to trace a predictive sequence, from physiological features to the growth response of species to elevated CO2 in pure culture, from the growth response in pure culture to competitive ability in pairwise mixtures and from pairwise competitive ability to shifts in the relative abundance of species in the full community of all six species. CO2 altered the dynamics of growth in a fashion consistent with known differences among major taxa in their ability to take up and use CO2. This pure-culture response was partly successful in predicting the outcome of competition in pairwise mixtures, especially the enhanced competitive ability of chlorophytes relative to cyanobacteria, although generally statistical support was weak. The competitive response in pairwise mixtures was a good predictor of changes in competitive ability in the full community. Hence, there is a potential for forging a logical chain of inferences for predicting how phytoplankton communities will respond to elevated CO2. Clearly further extensive experiments will be required to validate this approach in the greater complexity found in diverse communities and environments of natural systems.
Data from: Grass competition overwhelms effects of herbivores and precipitation on early tree establishment in Serengeti
1. Savanna ecosystems span a diverse range of climates, edaphic conditions and disturbance regimes, the complexity of which has stimulated long-standing interest in the mechanisms that maintain tree-grass coexistence. One hypothesis suggests that tree establishment is strongly limited by one or several demographic bottlenecks at early stages of the tree life cycle. A major impediment to testing this hypothesis is the lack of data on the relative strengths of different bottlenecks across key environmental gradients. 2. To identify demographic bottlenecks that limit early tree establishment (0-18 months), we conducted a series of transplant experiments with two savanna trees species (Acacia robusta and A. tortilis) across a natural rainfall and soil fertility gradient in the Serengeti ecosystem, Tanzania. We tested the interactive effects of precipitation, herbivory, seed scarification, grass competition, water limitation and tree species identity on two key life stages: germination and early seedling survival (0-2 months) and juvenile seedling survival (2-18 months). 3. Germination and early seedling survival increased as a function of rainfall, in the absence of herbivores and when seeds were scarified. Juvenile seedling survival, in contrast, decreased with rainfall but increased in the absence of herbivores. Grass removal had the single strongest (positive) effect on juvenile seedling survival of any treatment. Soil moisture monitoring and grass-addition treatments revealed that grasses negatively affected seedlings in ways that were not necessarily linked to soil moisture. 4. A demographic model combining all effects across early life stages showed that the strength of grass competition on juvenile seedling survival was the key factor limiting early tree establishment. While rainfall had an unexpected opposing effect on the two life stages, the net effect of mean annual precipitation on early tree establishment was positive. 5. Synthesis: Successful tree establishment in Serengeti is maximized by a seemingly unlikely sequence of events: (1) scarification of seeds by browsers, (2) heavy rainfall to promote germination, (3) intensive grazing (but absence of browsers) and (4) dry conditions during juvenile seedling growth (>2 months) to reduce competition with grasses. By considering a wide suite of conditions and their interactions, our experimental results are relevant to ongoing debates about savanna vegetation dynamics and structural shifts in tree:grass ratios.
The effect of inter-and intraspecific competition on individual and population niche widths – a four-decade study on two interacting salmonids
<p>Competition is assumed to shape niche widths, affecting species survival and coexistence. Expectedly, high interspecific competition will reduce population niche widths, whereas high intraspecific competition will do the opposite. Here we test in situ how intra- and interspecific competition affects trophic resource use and the individual and population niche widths of two lacustrine fish species, Arctic charr and brown trout, covering a 40 year study period with highly contrasting competitive impacts prior to and following a large-scale fish culling experiment. Initially, an overcrowded Arctic charr population dominated the study system, with brown trout being nearly absent. The culling experiment reduced the littoral Arctic charr density by 80%, whereupon brown trout gradually increased its density in the system. Thus, over the study period, the Arctic charr population went from high to low intraspecific competition, followed by increasing interspecific competition with brown trout. As hypothesized, the relaxed intraspecific competition following the experimental culling reduced individual diet specialization and compressed population niche width of Arctic charr. During the initial increase of the brown trout population, there was a large dietary overlap between the two species. Over the subsequent intensified interspecific competition from the population build-up of brown trout, their trophic niche overlap chiefly declined due to a dietary shift of Arctic charr towards enhanced zooplankton consumption. Contrary to theoretical expectations, the individual and population niche widths of Arctic charr increased with intensified interspecific competition. In contrast, the diet and niche width of brown trout remained stable over time, confirming its competitive superiority. The large-scale culling experiment and associated long-term research revealed pronounced temporal dynamics in trophic niche and resource use of the inferior competitor, substantiating that intra- and interspecific competition have large and contrasting impacts on individual and population niches.</p>
Data from: Isodars unveil asymmetric effects on habitat use caused by competition between two endangered species
In order for competing species to coexist, segregation on some ecological niche component is required and is often mediated by differential habitat use. When unequal competitors are involved, the dominant species tends to displace the subordinate one to its less preferred habitat. Here, we use habitat isodars, an approach which reflects evolutionary stable strategies of habitat selection, to evaluate whether interspecific competition between two competing species with distinct habitat preferences, the little bustard Tetrax tetrax and the great bustard Otis tarda, modulates their habitat use. Field data on these endangered species demonstrate that unequal competitors can coexist without completely segregating on their preferred habitats. The negatively sloped isodar of the subordinate little bustard unveils its competition with the dominant great bustard. Interference from great bustards in secondary cereal habitats reinforces use of preferred natural habitat by little bustards. Studies of density-dependent habitat selection by a single-species can thus aid in identifying the effects of competition on community composition, and guide the conservation of at-risk species. Isodars, in particular, represent a promising method to gain clear knowledge on interspecific competition for species in which experimental manipulations are not feasible.
The effect of urbanization and temperature on thermal tolerance, foraging performance, and competition in cavity dwelling ants
<p>Human disturbance including rapid urbanization and increased temperatures can have profound effects on the ecology of local populations. Eusocial insects, such as ants, have adapted to stressors of increasing temperature and urbanization, however these evolutionary responses are not consistent among populations across geographic space. Here we asked how urbanization and incubation temperature influence critical thermal maximum (CT<sub>max</sub>) and various ecologically relevant behaviors in three ant species in urban and rural locations in Worcester, MA, USA. We did this by incubating colonies of three species of cavity dwelling ant (<em>Aphaenogaster picea, Tapinoma sessile, </em>and <em>Temnothorax longispinosus</em>) from 2 habitat types (Rural and Urban), for 60-days at multiple temperatures. We found that incubation temperature, urbanization and species of ant all significantly affected overall colony critical thermal maximum. We also found that recruitment time, colonization time and defense response were significantly affected by incubation temperature and varied between species of ant, while recruitment and colonization time were additionally affected by urbanization. These variable changes in performance and competitive traits across species suggest that responses to urbanization and shifting temperatures are not universal across species. Changes in behavioral responses caused by urbanization may disrupt biodiversity, creating unusual competitive environments as a consequence of natural adaptations and cause both direct and indirect mechanisms for which human disturbance can lead to local species extinction.</p> <p> </p>
Effect of heat shock and competition on the antioxidant system of round goby
<p><span><span><span><span><span><span><span><span><span><span><span>Monitoring oxidative stress biomarkers has become a powerful and common tool to estimate organismal condition and response to endogenous and environmental factors. In the present study, we used round goby (<i>Neogobius melanostomus</i>) from non-native European populations, as a model species to test sex differences in oxidative stress biomarkers. Considering sex differences in reproductive investment, we hypothesized that males would display lower resistance to abiotic stress. Fish were exposed to a heat shock (temperature elevated by 10°C) for 1h, 6h, and 12h and catalase activity (CAT), reduced glutathione (GSH), total antioxidant capacity (TAC) and lipid peroxidation (LPO) were measured in liver and muscle tissues. Liver of males was significantly more responsive compared to liver of females in all tested parameters. GSH was found to be the most responsive to heat stress exposure in both sexes. The results supported our hypothesis that male reproductive investment (territoriality, courtship, and brood care) and likelihood of only a single spawning period in their lifetime influenced on higher sensitivity of their antioxidant defence. On the other hand, for females antioxidant defence is considered more important to survive the environmental changes and successfully reproduce in the next season. Our experiments exposed fish to acute thermal stress. Further research should determine the effects of exposure to chronic thermal stress to corroborate our understanding on sex differences in antioxidant defence in the round goby.</span></span></span></span></span></span></span></span></span></span></span></p>
Data from: A generalist nematode destabilizes plant competition: no evidence for direct effects, but strong evidence for indirect effects on rhizobia abundance
<p><span>1. Difficulties quantifying pathogen load and mutualist abundance limit our ability to connect disease dynamics to host community ecology. For example, specific predictions about how differential pathogen load is hypothesized to drive host competitive outcomes are rarely tested. Additionally, although infection is known to affect mutualists, we rarely measure the magnitude of pathogen effects on mutualist abundance across host competitive contexts. <span>We </span>tested for both mechanisms <span>in </span>a plant-rhizobia-nematode system.</span></p> <p><span><span>2. We </span>paired the legume <i>Medicago lupulina </i>with intra- and interspecific plant <span>com</span>petitors, with and without a generalist nematode parasite <i>Meloidogyne </i>sp. <span>Relative </span>change in plant biomass was used to determine how nematode inoculation <span>affected </span>plant competitive outcomes. <span>We</span> counted nematode galls to test for direct effects of parasitism on plant competition, and rhizobia nodules to test for indirect effects <span>of </span>nematode presence on rhizobia abundance.</span></p> <p>3. Parasites were destabilizing despite similar nematode load across competition treatments. During inter- compared to intraspecific competition, nematode <span>inoculation </span>decreased nodulation on <i>M. lupulina</i>, increased nodulation on <i>Trifolium repens</i>, and had no effect on nodulation on <i>Chamaecrista fasciculata.</i></p> <p><span><span>4. We </span>found no support for hypothesized direct effects of nematode load on competitive outcomes, and strong but idiosyncratic indirect effects of nematode <span>inoculation </span>on rhizobia abundance.</span></p>
Disentangling the effects of biomass and productivity in plant competition
<p>The relationship between competition and productivity in plant communities is unclear, likely due to (i) a confusion in the literature between productivity and biomass, (ii) the lack of studies assessing variation in competition in all combinations of biomass and productivity. We assessed the outcome of plant-plant interactions by removing neighbors around five focal species in 14 herbaceous communities with contrasting biomasses and productivities: meadows with high biomass and productivity, heathlands with high biomass and low productivity, understorey communities of deciduous forests with low biomass and high productivity and calcareous grasslands with low biomass and low productivity. Competition intensity was quantified with the relative interaction index (RII) calculated for both survival and growth of the transplanted targets assessed with the increase in leaf number. To examine which traits better explain variation in competition and what drives variation in diversity, we e also quantified litter decomposition rate, species composition and diversity and six morphological traits related to plant size and growth rate for eight dominant species of each community. Our main questions were: (i) is competition mostly related to biomass or productivity? (ii) which traits of the community dominants better explain variation in competition? (iii) is variation in competition and related-traits correlated with variation in diversity? Competition for survival significantly increased with increasing community biomass (but not productivity). In addition, competition for survival increased with the size traits and competitive effects of the dominant species of the communities, while diversity decreased. Competition for growth also increased with increasing productivity, but only for high biomass communities. Additionally, the increase in competition for growth with increasing soil fertility, as measured with litter decomposition rate, was only due to an increase in target growth in plots without neighbors and was unrelated to community competitive effects and species diversity. The results of our study illustrate how the confusion between productivity and biomass could have contributed to the longstanding debate on variation in competition along productivity gradients and its consequence for diversity.</p>
Data from: Climate and competition effects on tree growth in Rocky Mountain forests
1. Climate is widely assumed to influence physiological and demographic processes in trees, and hence forest composition, biomass and range limits. Growth in trees is an important barometer of climate change impacts on forests as growth is highly correlated with other demographic processes including tree mortality and fecundity. 2. We investigated the main drivers of diameter growth for five common tree species occurring in the Rocky Mountains of the western United States using non-linear regression methods. We quantified growth at the individual tree level from tree core samples collected across broad environmental gradients. We estimated the effects of both climate variation and biotic interactions on growth processes and tested for evidence that disjunct populations of a species respond differentially to climate. 3. Relationships between tree growth and climate varied by species and location. Growth in all species responded positively to increases in annual moisture up to a threshold level. Modest linear responses to temperature, both positive and negative, were observed at many sites. However, model results also revealed evidence for differentiated responses to local site conditions in all species. In severe environments in particular, growth responses varied non-linearly with temperature. For example, in northerly cold locations pronounced positive growth responses to increasing temperatures were observed. In warmer southerly climates, growth responses were unimodal, declining markedly above a threshold temperature level. 4. Net effects from biotic interactions on diameter growth were negative for all study species. Evidence for facilitative effects was not detected. For some species, competitive effects more strongly influenced growth performance than climate. Competitive interactions also modified growth responses to climate to some degree. 6. Synthesis. These analyses suggest that climate change will have complex, species specific effects on tree growth in the Rocky Mountains due to non-linear responses to climate, differentiated growth processes that vary by location and complex species interactions that impact growth and potentially modify responses to climate. Thus, robust model simulations of future growth responses to climate trends may need to integrate realistic scenarios of neighborhood effects as well as variability in tree performance attributed to differentiated populations.
Data from: A rapid and cost-effective quantitative microsatellite genotyping protocol to estimate intraspecific competition in protist microcosm experiments
High levels of intra-specific variation are commonly observed in natural microbial populations, yet the consequences of this variation for ecological and evolutionary processes remains poorly understood. Protists are excellent experimental models for investigating fundamental and applied questions in ecology and evolution, but studying intra-specific variation remains a challenge due to a lack of molecular resources to aid in quantifying and distinguishing strains during experiments. Here we present a molecular method, quantitative microsatellite genotyping, to accurately quantify strain specific frequencies from microcosm experiments of the marine flagellate Oxyrrhis marina, both between many pairs of strains and between strains in a multi-strain mixture. We find that for pairs of strains the method is effective for relative frequencies as low as 0.02 and with around 99% accuracy. The method is able to quantify four strains reasonably well, though less accurately than for pairs (range 92%-97% accuracy). This makes accessible a cheap and easy to implement method for quantifying strain (or allele) frequencies, and is suitable for use in a broad range of single celled eukaryotes (Protists) where copy number should correlate well with number of individuals (i.e. cells). This opens up the possibility of examining the role of intra-specific variation using experimental protist microcosms.
Male age alone predicts paternity success under sperm competition when effects of age and past mating effort are experimentally separated
<p>Older males often perform poorly under post-copulatory sexual selection. It is unclear, however, whether reproductive senescence is due to male age itself or the accumulated costs of the higher lifetime mating effort that is usually associated with male age. To date, very few studies have accounted for mating history and sperm storage when testing the effect of male age on sperm traits, and none test how age and past mating history influence paternity success under sperm competition. Here, we experimentally manipulate male mating history to tease apart its effects from that of age on ejaculate traits and paternity in the mosquitofish<i>, Gambusia holbrooki</i>. We found that old, naive males had more sperm than old, experienced males, while the reverse was true for young males. In contrast, neither male age nor mating history affected sperm velocity. Finally, using artificial insemination to experimentally control the number of sperm per male, we found that old males sired significantly more offspring than young males independently of their mating history. Our results highlight that the general pattern of male reproductive senescence described in many taxa may often be affected by two naturally confounding factors, male mating history and sperm age, rather than male age itself.</p>
Competitive effects of a dominant palm on sapling performance in a Neotropical rainforest
<p>Eradication of herbivores, due to human disturbances, produces a demographic outburst of highly competitive prey species, which in turn reduces plant species diversity. This happens at Los Tuxtlas tropical rainforest, Mexico, where a population outburst of the understory palm <em>Astrocaryum mexicanum</em> is ostensibly excluding tree species, but how this is occurring is still unknown. We used a neighborhood approach to explore the effects of palm shading and palm crowding on the survival and growth (RGR) of saplings of six common tree species. Sixteen to 32 saplings (1.5-2.5 m height) per species were used as focal individuals of circular neighborhoods (4 m radius), which included palms ≥ 1cm stem length potentially competing for light or soil resources. Shading was estimated using hemispherical photographs. Overall, survivorship was high combining all species (93.8%/yr). In most species shading produced a displacement of the crown, which increased with specific leaf area of species. In three species shading had a negative effect on RGR without any effect of crowding, the contrary occurred in two species, and in one species no effects were found. No effects due to trees (DBH ≥ 1cm) crowding were detected. The shading effect increased with species leaf dry matter content (LDMC), while the effect of crowding declined with LDMC and increased with sapling total leaf area. We argue that the species-dependent palm shading/crowding effects were related to the shade tolerance of sapling species. In the long-term such species-specific responses could have consequences for forest structure and composition, as saplings develop to mature stages.</p>
Effect of defoliation on competition between Poa annua plants
<p>This dataset shows shoot and root masses of Poa annua plants grown in split root boxes with N supplied equally or unequally to the two chambers. In expt 1, half the plants are subject to defoliation, while in expt 2, pairs of plants are grown in a three-chamber split root box with a shared center (competition) chamber. N was supplied either to the outer or center chamber, and half of the "competitor" plants were subject to defoliation</p>
Subannual phenology and the effect of staggered fruit ripening on dispersal competition
<p>Seed dispersal mutualisms evolve in complex communities of plants and frugivorous animals, within which indirect interactions such as competition and facilitation can occur. Many tropical plants reproduce subannually in multiple episodes per year. Yet, the consequences of episodic reproduction on interactions with seed dispersers remains largely unexplored. We studied <em>Guarea guidonia</em> (<em>Meliaceae</em>), a subannually reproducting tree, to examine temporal variation in seed dispersal within a tropical forested landscape in the central Dominican Republic. We hypothesized that foraging by seed dispersers would (a) increase with daily ripe fruit set on focal trees, (b) decrease with increasing ripe fruit biomass of neighboring plants, and (c) decrease in response to the fruiting periods of other taxa at the landscape scale. Over 18 months, we tracked the phenology of 24 focal trees and quantified foraging during fruiting phases through repeated observations, simultenously measuring seed dispersal in traps beneath isolated perches across the study landscape. Date was the only clear predictor of frugivore visitation, with early and late peaks in activity during the 5-month fruiting period. The midseason decline in foraging at focal trees matched a decline in <em>Guarea</em> dispersal to seed traps independently of fruit abundance. Declines in foraging associated with <em>Guarea</em> were inversely related to peak dispersal of higher quality lipid-rich fruiting species. Our results suggest that multiple flowering episodes and subsequent asynchronous fruit ripening of low-quality fruits can reduce competititive pressure from other high quality fruiting species, implying that this potentialy bet-hedging strategy may be an overlooked factor in the evolution of subannual reproduction.</p>
The effects of novel leaf litter deposition on competitive, predator–prey and host–parasite interactions of American toad larvae
<p>Wetland plant communities are changing rapidly due to a wide range of human activities. The deposition of leaf litter from novel plant communities can alter both the chemical and physical habitat of aquatic ecosystems. Lesser understood are the ecological consequences of novel leaf litter inputs in aquatic communities. Towards this goal, we used two plant invasion scenarios (comparing native black huckleberry to exotic autumn olive and native swamp loosestrife to exotic purple loosestrife) to simulate a shift in wetland plant communities. In this study, we investigated the effects of novel leaf litter leachates on three aquatic ecological interactions: intraspecific competition, predation, and parasitism. We examined how leaf litter leachates influence the interactions of American toad larvae (Anaxyrus americanus) with their conspecifics, a dragonfly predator (Anax spp.) and a trematode parasite (Echinostomatidae). We found that leaf litter type influenced competitive interactions only for the huckleberry versus autumn olive comparison. We did not detect any effects of leaf litter type on predator-prey interactions. Finally, litter type strongly influenced host-parasite interactions for both leaf litter comparisons, altering host susceptibility, parasite survival and net infection rates. These results highlight the breadth of potential ecological repercussions of shifting wetland plant communities for native ecosystems.</p>
Effect of Two Preventive Exercise Programs for Swimmer's Shoulder on the Torque of Shoulder Rotator Muscles in Competitive Swimmers
ClinicalTrials.gov study NCT06552585. IPD Sharing: NO. Countries: 1. Publications: 1.
The Effects of Omega-3 Status and Supplementation on Tendon Structure in Competitive Athletes
ClinicalTrials.gov study NCT03880149. IPD Sharing: Not stated. Countries: 1. Publications: 4.
Effects of a Mindfulness Programme on Competitive Athletes With Physical Disabilities
ClinicalTrials.gov study NCT06184217. IPD Sharing: NO. Countries: 1. Publications: 5.
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Allen Brain Atlas
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Annotated Behaviour and Observability Dataset (ABODe)
ABODe is a University of Edinburgh DataShare dataset for behavior classification in group-housed mice using home-cage video, identities, bounding boxes, ground-plate positions, and annotator labels.
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