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Abstract
Over half a century ago, British developmental biologist Conrad Hal Waddington proposed the idea of canalization, that is, homeostasis in development. Since the breakthrough that was made by Rutherford and Lindquist (1998), who proposed a role of Hsp90 in developmental buffering, chaperones have gained much attention in the study of canalization. However, recent studies have revealed that a number of other molecules are also potentially involved in canalization. Here, I introduce the emerging role of DnaJ chaperones in canalization. I also discuss how the expression levels of such buffering molecules can be altered, thereby altering organismal development. Since developmental robustness is maternally inherited in various organisms, I propose that dynamic bet hedging, an increase in within-clutch variation in offspring phenotypes that is caused by unpredictable environmental challenges to the mothers, plays a key role in altering the expression levels of buffering molecules. Investigating dynamic bet hedging at the molecular level and how it impacts upon morphological phenotypes will help our understanding of the molecular mechanisms of canalization and evolutionary processes.
Highlights
Chaperones have been studied as important players in the maintenance of homeostasis
When mRNAs are translated, some of the nascent peptides are folded by co-translational folding at the ribosomes, while others are transferred to the endoplasmic reticulum (ER) for correct folding [1]
Recent studies have revealed a wide range of molecules that are involved in canalization, including DnaJ chaperones
Summary
Chaperones have been studied as important players in the maintenance of homeostasis. They are involved from the birth to the death of proteins. Upon environmental stress or excess production of nascent peptide chains, some peptides are refolded and they escape from aggregation, while others are delivered to the ER for degradation None of these processes are conducted without chaperones. In his book ‘The Strategy of the Genes’, Conrad Hal Waddington originally coined the term ‘canalization’ or ‘homeorhesis’, which differs from ‘physiological homeostasis’, referring to some physiological state that is being held constant, and ‘genetic homeostasis’ in which the constant feature of the system is its set of gene frequencies [11] (p.44) He understood the developmental process as a ball falling down a canalized slope in a metaphoric multi-dimensional phase space called the ‘epigenetic landscape’ underpinned by ‘a complex system’ [11]. As an immense amount of work has been published on chaperones in relation to evolution (a topic covered by recent reviews, including [14,15] and references therein), in this review I focus on discussing how studies on the role of chaperones in developmental robustness and evolution can move forward by integration with ecological studies, leading to a better understanding of the interaction between the environment and the evolution of animal forms
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