Taste buds on the anterior part of the tongue develop in conjunction with epithelial-mesenchymal specializations in the form of gustatory (taste) papillae. Sonic hedgehog (Shh) and Bone Morphogenetic Protein 4 (BMP4) are expressed in developing taste papillae, but the roles of these signaling molecules in specification of taste bud progenitors and in papillary morphogenesis are unclear. We show here that BMP4 is not expressed in the early tongue, but is precisely coexpressed with Shh in papillary placodes, which serve as a signaling center for both gustatory and papillary development. To elucidate the role of Shh, we used an in vitro model of mouse fungiform papillary development to determine the effects of two functional inhibitors of Shh signaling: anti-Shh (5E1) antibody and cyclopamine. Cultured E11.5 tongue explants express Shh and BMP4(LacZ) in a pattern similar to that of intact embryos, localizing to developing papillary placodes after 2 days in culture. Tongues cultured with 5E1 antibody continue to express these genes in papillary patterns but develop more papillae that are larger and closer together than in controls. Tongues cultured with cyclopamine have a dose-dependent expansion of Shh and BMP4(LacZ) expression domains. Both antibody-treated and cyclopamine-treated tongue explants also are smaller than controls. Taken together, these results suggest that, although Shh is not involved in the initial specification of papillary placodes, Shh does play two key roles during pmcry development: (1) as a morphogen that directs cells toward a nonpapillary fate, and (2) as a mitogen, causing expansion of the interplacodal epithelium and underlying mesenchyme.
Research on the thermal ecology and physiology of free‐living organisms is accelerating as scientists and managers recognize the urgency of the global biodiversity crisis brought on by climate change. As ectotherms, temperature fundamentally affects most aspects of the lives of amphibians and reptiles, making them excellent models for studying how animals are impacted by changing temperatures. As research on this group of organisms accelerates, it is essential to maintain consistent and optimal methodology so that results can be compared across groups and over time. This review addresses the utility of reptiles and amphibians as model organisms for thermal studies by reviewing the best practices for research on their thermal ecology and physiology, and by highlighting key studies that have advanced the field with new and improved methods. We end by presenting several areas where reptiles and amphibians show great promise for further advancing our understanding of how temperature relations between organisms and their environments are impacted by global climate change.
Effects of global change (i.e. urbanization, climate change) on adult organisms are readily used to predict the persistence of populations. However, effects on embryo survival and patterns of development are less studied, even though embryos are particularly sensitive to abiotic conditions that are altered by global change (e.g. temperature). In reptiles, relatively warm incubation temperatures increase developmental rate and often enhance fitness-relevant phenotypes, but extremely high temperatures cause death. Due to the urban heat island effect, human-altered habitats (i.e. cities) potentially create unusually warm nest conditions that differ from adjacent natural areas in both mean and extreme temperatures. Such variation may exert selection pressures on embryos. To address this, we measured soil temperatures in places where the Puerto Rican crested anole lizard () nests in both city and forest habitats. We bred anoles in the laboratory and subjected their eggs to five incubation treatments that mimicked temperature regimes from the field, three of which included brief exposure to extremely high temperatures (i.e. thermal spikes) measured in the city. We monitored growth and survival of hatchlings in the laboratory for 3 months and found that warmer, city temperatures increase developmental rate, but brief, thermal spikes reduce survival. Hatchling growth and survival were unaffected by incubation treatment. The urban landscape can potentially create selection pressures that influence organisms at early (e.g. embryo) and late life stages. Thus, research aimed at quantifying the impacts of urbanization on wildlife populations must include multiple life stages to gain a comprehensive understanding of this important aspect of global change.
Most studies of thermal tolerance use adults, but early-life stages (e.g. embryos) are often more sensitive to thermal agitation. Studies that examine effects on embryos rarely assess the potential for thermal tolerance to change with ontogeny or how effects differ among sympatric species, and often utilize unrealistic temperature treatments. We used thermal fluctuations from nests within the urbanheat island to determine how thermal tolerance of embryos changes across development and differs among two sympatric lizard species (Anolis sagrei and Anolis cristatellus). We applied fluctuations that varied in frequency and magnitude at different times during development and measured effects on embryo physiology and survival, and hatchling morphology, growth and survival. Thermal tolerance differed between the species by ∼2°C: embryos of A. sagrei, a lizard that prefers warmer, open-canopy microhabitats, were more robust to thermal stress than embryos of A. cristatellus, which prefers cooler, closed-canopy microhabitats. Moreover, thermal tolerance changed through development; however, the nature of this change differed between the species. For A. cristatellus, thermal tolerance was greatest mid-development. For A. sagrei, the relationship was not statistically clear. The greatest effects of thermal stress were on embryo and hatchling survival and embryo physiology. Hatchling morphology and growth were less affected. Inter-specific responses and the timing of stochastic thermal events with respect to development have important effects on embryo mortality. Thus, research that integrates ecologically meaningful thermal treatments, considers multiple life-history stages and examines interspecific responses will be critical to make robust predictions of the impacts of global change on wildlife.
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