Species diverge eco-morphologically through the continuous action of natural selection on functionally important structures, producing alternative adaptive morphologies. In cichlid fishes, the oral and pharyngeal jaws are such key structures. Adaptive variation in jaw morphology contributes to trophic specialization, which is hypothesized to fuel their rapid speciation in the East African Great Lakes. Much is known about the genes involved in cichlid jaw and craniofacial development. However, it is still unclear what salient sources of variation gave rise to trophic-niche specialization, facilitating adaptive radiation. Here, we explore two sources of transcriptional variation that may underlie species-specific disparities in jaw morphology. Using whole transcriptome RNA-sequencing, we analyze differences in gene expression and alternative splicing, at the end of postlarval development, in fully functional jaws of six species of cichlids from the Lake Tanganyika tribe Tropheini. Our data reveal a surprisingly high degree of alternative splicing events compared with gene expression differences among species and trophic types. This suggests that differential trophic adaptation of the jaw apparatus may have been shaped by transcriptional rewiring of splicing as well as gene expression variation during the rapid radiation of the Tropheini. Specifically, genes undergoing splicing across most species were found to be enriched for pharyngeal jaw gene ontology terms. Overall, jaw transcriptional patterns at postlarval developmental stage were highly dynamic and species-specific. In conclusion, this work indicates that shifts in alternative splicing could have played a more important role in cichlid adaptive radiation, and possibly adaptive radiation in general, than currently recognized.
Whether an animal is trying to escape from a predator, avoid a fall or perform a more mundane task, the effectiveness of its sensory feedback is constrained by sensorimotor delays. Here, we combine electrophysiological experiments, systematic reviews of the literature and biophysical models to determine how delays associated with the fastest locomotor reflex scale with size in terrestrial mammals. Nerve conduction delay is one contributor, and increases strongly with animal size. Sensing, synaptic and neuromuscular junction delays also contribute, and we approximate each as a constant value independent of animal size. Muscle's electromechanical and force generation delays increase more moderately with animal size than nerve conduction delay, but their total contribution exceeds that of the four neural delays. The sum of these six component delays, termed total delay, increases with animal size in proportion to -large mammals experience total delays 17 times longer than small mammals. The slower movement times of large animals mostly offset their long delays resulting in a more modest, but perhaps still significant, doubling of their total delay relative to movement duration when compared with their smaller counterparts. Irrespective of size, sensorimotor delay is likely a challenge for all mammals, particularly during fast running.
Sensorimotor control is greatly affected by two factors-the time it takes for an animal to sense and respond to stimuli (responsiveness), and the ability of an animal to distinguish between sensory stimuli and generate graded muscle forces (resolution). Here, we demonstrate that anatomical limitations force a necessary trade-off between responsiveness and resolution with increases in animal size. To determine whether responsiveness is prioritized over resolution, or resolution over responsiveness, we studied how size influences the physiological mechanisms underlying sensorimotor control. Using both new electrophysiological experiments and existing data, we determined the maximum axonal conduction velocity (CV) in animals ranging in size from shrews to elephants. Over the 100-fold increase in leg length, CV was nearly constant, increasing proportionally with mass to the 0.04 power. As a consequence, larger animals are burdened with relatively long physiological delays, which may have broad implications for their behaviour, ecology and evolution, including constraining agility and requiring prediction to help control movements.
Sensorimotor control is greatly affected by two factors-the time it takes for an animal to sense and respond to stimuli (responsiveness), and the ability of an animal to distinguish between sensory stimuli and generate graded muscle forces (resolution). Here, we demonstrate that anatomical limitations force a necessary trade-off between responsiveness and resolution with increases in animal size. To determine whether responsiveness is prioritized over resolution, or resolution over responsiveness, we studied how size influences the physiological mechanisms underlying sensorimotor control. Using both new electrophysiological experiments and existing data, we determined the maximum axonal conduction velocity (CV) in animals ranging in size from shrews to elephants. Over the 100-fold increase in leg length, CV was nearly constant, increasing proportionally with mass to the 0.04 power. As a consequence, larger animals are burdened with relatively long physiological delays, which may have broad implications for their behaviour, ecology and evolution, including constraining agility and requiring prediction to help control movements.
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