Developmental systems integrate inputs of variation from different origins into the observable variation of the resulting phenotype. Different components of phenotypic variation can be distinguished that correspond to those inputs, but the response of the system to each input factor can be modulated by other factors, so that there are interactions among the factors. This paper uses the concept of the target phenotype, introduced by Nijhout and Davidowitz to designate the expected phenotype for a given genotype and environment, to provide definitions and explanations of key concepts. This logic is put to use to explore the factors contributing to fluctuating asymmetry. To illustrate how genetic variation, phenotypic plasticity in response to temperature variation, and random developmental noise interact, this study uses computer simulations based on a simplified developmental model of a trait. The simulations show extensive interactions among the input factors, which can be explained by the non-linear nature of the developmental model. As a result, all the loci that control the developmental parameters end up affecting the resulting phenotypic trait, its reaction norm in response to temperature changes, and also its fluctuating asymmetry. Further, temperature (or possibly other environmental factors) can affect both trait values and the amount of fluctuating asymmetry without any involvement of stress. The model is broadly consistent with what is known from actual biological systems, and the results obtained from it have far-reaching implications for interpreting observations of the different types of phenotypic variation.
The primary function of leaves is to provide an interface between plants and their environment for gas exchange, light exposure and thermoregulation. Leaves have, therefore a central contribution to plant fitness by allowing an efficient absorption of sunlight energy through photosynthesis to ensure an optimal growth. Their final geometry will result from a balance between the need to maximize energy uptake while minimizing the damage caused by environmental stresses. This intimate relationship between leaf and its surroundings has led to an enormous diversification in leaf forms. Leaf shape varies between species, populations, individuals or even within identical genotypes when those are subjected to different environmental conditions. For instance, the extent of leaf margin dissection has, for long, been found to inversely correlate with the mean annual temperature, such that Paleobotanists have used models based on leaf shape to predict the paleoclimate from fossil flora. Leaf growth is not only dependent on temperature but is also regulated by many other environmental factors such as light quality and intensity or ambient humidity. This raises the question of how the different signals can be integrated at the molecular level and converted into clear developmental decisions. Several recent studies have started to shed the light on the molecular mechanisms that connect the environmental sensing with organ-growth and patterning. In this review, we discuss the current knowledge on the influence of different environmental signals on leaf size and shape, their integration as well as their importance for plant adaptation.