Abstract:Polyploidy in combination with parthenogenesis offers advantages for plasticity and the evolution of a broad ecological tolerance of species. Therefore, a positive correlation between the level of ploidy and increasing latitude as a surrogate for environmental harshness has been suggested. Such a positive correlation is well documented for plants, but examples for animals are still rare. Species of flatworms (Platyhelminthes) are widely distributed, show a remarkably wide range of chromosome numbers, and offer… Show more
“…These orders exhibit a significant difference in genome size (and hence probably cell size) [ t ‐test: P = 1.005 × 10 −7 ; calculated from data in Gregory, 2018; see also Wyngaard & Rasch, 2000]. Similarly, aquatic species living in cold environments tend to have larger genomes (Dufresne & Jeffery, 2011; Lorch et al ., 2016; Alfsnes et al ., 2017; Jeffery, Yampolsky, & Gregory, 2017), suggesting a cell‐size parallel with Bergmann's rule, at least for aquatic animals. In general, animals increase body size mainly through cell proliferation during early development, but by cell growth in later life (Kammenga et al ., 2007; Czarnołęski et al ., 2008, but see Aguilar‐Alberola & Mesquita‐Joanes, 2014; Horne et al ., 2019) and this fits with the TSR being less pronounced for egg size, manifesting itself in later life stages (Forster, Hirst, & Atkinson, 2011; Forster & Hirst, 2012).…”
Section: The Dependency Of T–s Responses On Growth and Developmentmentioning
Body size is central to ecology at levels ranging from organismal fecundity to the functioning of communities and ecosystems. Understanding temperature‐induced variations in body size is therefore of fundamental and applied interest, yet thermal responses of body size remain poorly understood. Temperature–size (T–S) responses tend to be negative (e.g. smaller body size at maturity when reared under warmer conditions), which has been termed the temperature–size rule (TSR). Explanations emphasize either physiological mechanisms (e.g. limitation of oxygen or other resources and temperature‐dependent resource allocation) or the adaptive value of either a large body size (e.g. to increase fecundity) or a short development time (e.g. in response to increased mortality in warm conditions). Oxygen limitation could act as a proximate factor, but we suggest it more likely constitutes a selective pressure to reduce body size in the warm: risks of oxygen limitation will be reduced as a consequence of evolution eliminating genotypes more prone to oxygen limitation. Thus, T–S responses can be explained by the ‘Ghost of Oxygen‐limitation Past’, whereby the resulting (evolved) T–S responses safeguard sufficient oxygen provisioning under warmer conditions, reflecting the balance between oxygen supply and demands experienced by ancestors. T–S responses vary considerably across species, but some of this variation is predictable. Body‐size reductions with warming are stronger in aquatic taxa than in terrestrial taxa. We discuss whether larger aquatic taxa may especially face greater risks of oxygen limitation as they grow, which may be manifested at the cellular level, the level of the gills and the whole‐organism level. In contrast to aquatic species, terrestrial ectotherms may be less prone to oxygen limitation and prioritize early maturity over large size, likely because overwintering is more challenging, with concomitant stronger end‐of season time constraints. Mechanisms related to time constraints and oxygen limitation are not mutually exclusive explanations for the TSR. Rather, these and other mechanisms may operate in tandem. But their relative importance may vary depending on the ecology and physiology of the species in question, explaining not only the general tendency of negative T–S responses but also variation in T–S responses among animals differing in mode of respiration (e.g. water breathers versus air breathers), genome size, voltinism and thermally associated behaviour (e.g. heliotherms).
“…These orders exhibit a significant difference in genome size (and hence probably cell size) [ t ‐test: P = 1.005 × 10 −7 ; calculated from data in Gregory, 2018; see also Wyngaard & Rasch, 2000]. Similarly, aquatic species living in cold environments tend to have larger genomes (Dufresne & Jeffery, 2011; Lorch et al ., 2016; Alfsnes et al ., 2017; Jeffery, Yampolsky, & Gregory, 2017), suggesting a cell‐size parallel with Bergmann's rule, at least for aquatic animals. In general, animals increase body size mainly through cell proliferation during early development, but by cell growth in later life (Kammenga et al ., 2007; Czarnołęski et al ., 2008, but see Aguilar‐Alberola & Mesquita‐Joanes, 2014; Horne et al ., 2019) and this fits with the TSR being less pronounced for egg size, manifesting itself in later life stages (Forster, Hirst, & Atkinson, 2011; Forster & Hirst, 2012).…”
Section: The Dependency Of T–s Responses On Growth and Developmentmentioning
Body size is central to ecology at levels ranging from organismal fecundity to the functioning of communities and ecosystems. Understanding temperature‐induced variations in body size is therefore of fundamental and applied interest, yet thermal responses of body size remain poorly understood. Temperature–size (T–S) responses tend to be negative (e.g. smaller body size at maturity when reared under warmer conditions), which has been termed the temperature–size rule (TSR). Explanations emphasize either physiological mechanisms (e.g. limitation of oxygen or other resources and temperature‐dependent resource allocation) or the adaptive value of either a large body size (e.g. to increase fecundity) or a short development time (e.g. in response to increased mortality in warm conditions). Oxygen limitation could act as a proximate factor, but we suggest it more likely constitutes a selective pressure to reduce body size in the warm: risks of oxygen limitation will be reduced as a consequence of evolution eliminating genotypes more prone to oxygen limitation. Thus, T–S responses can be explained by the ‘Ghost of Oxygen‐limitation Past’, whereby the resulting (evolved) T–S responses safeguard sufficient oxygen provisioning under warmer conditions, reflecting the balance between oxygen supply and demands experienced by ancestors. T–S responses vary considerably across species, but some of this variation is predictable. Body‐size reductions with warming are stronger in aquatic taxa than in terrestrial taxa. We discuss whether larger aquatic taxa may especially face greater risks of oxygen limitation as they grow, which may be manifested at the cellular level, the level of the gills and the whole‐organism level. In contrast to aquatic species, terrestrial ectotherms may be less prone to oxygen limitation and prioritize early maturity over large size, likely because overwintering is more challenging, with concomitant stronger end‐of season time constraints. Mechanisms related to time constraints and oxygen limitation are not mutually exclusive explanations for the TSR. Rather, these and other mechanisms may operate in tandem. But their relative importance may vary depending on the ecology and physiology of the species in question, explaining not only the general tendency of negative T–S responses but also variation in T–S responses among animals differing in mode of respiration (e.g. water breathers versus air breathers), genome size, voltinism and thermally associated behaviour (e.g. heliotherms).
“…Indeed, as more flatworm species are incorporated into molecular studies, a greater number of genomic peculiarities are being discovered. Examples include several instances of polyploidy [16], high degrees of genomic polymorphy [17], [18] and 18S rDNA polymorphisms [19]. In the case of mitochondrial (mt) genomes, this variation is well illustrated by technical difficulties in protocols that are relatively straightforward with other taxa.…”
The first three mitochondrial (mt) genomes of endosymbiotic turbellarian flatworms are characterised for the rhabdocoels Graffilla buccinicola, Syndesmis echinorum and S. kurakaikina. Interspecific comparison of the three newly obtained sequences and the only previously characterised rhabdocoel, the free-living species Bothromesostoma personatum, reveals high mt genomic variability, including numerous rearrangements. The first intrageneric comparison within rhabdocoels shows that gene order is not fully conserved even at species levels. Atp8, until recently assumed absent in flatworms, was putatively annotated in two sequences. Selection pressure was tested in a phylogenetic framework and is shown to be significantly relaxed in this and another protein-coding gene: cox1. If present, atp8 appears highly derived in platyhelminths and its functionality needs to be addressed in future research. Our findings for the first time allude to a large degree of undiscovered (mt) genomic plasticity in rhabdocoels. It merits further attention whether this variation is correlated with a symbiotic lifestyle. Our results illustrate that this phenomenon is widespread in flatworms as a whole and not exclusive to the better-studied neodermatans.
“…In studies on the ecophysiology of ectotherms, a field gaining more and more interest deals with the regulation and consequences of cell size [1]. For ectotherms, patterns in cell size across thermal clines associated with latitude and altitude have been documented, where animals are generally composed of larger cells in the cold [2][3][4]. In addition, when ambient temperatures are experimentally lowered while rearing ectotherms, cell size tends to increase and this holds for phyla as diverse as nematodes (Caenorhabditis elegans; [5]), arthropods (Daphnia magna; [6]) and chordates (the edible frog Pelophylax esculentus; [7]).…”
There is renewed interest in the regulation and consequences of cell size adaptations in studies on understanding the ecophysiology of ectotherms. Here we test if induction of triploidy, which increases cell size in zebrafish (Danio rerio), makes for a good model system to study consequences of cell size. Ideally, diploid and triploid zebrafish should differ in cell size, but should otherwise be comparable in order to be suitable as a model. We induced triploidy by cold shock and compared diploid and triploid zebrafish larvae under standard rearing conditions for differences in genome size, cell size and cell number, development, growth and swimming performance and expression of housekeeping genes and hsp70.1. Triploid zebrafish have larger but fewer cells, and the increase in cell size matched the increase in genome size (+ 50%). Under standard conditions, patterns in gene expression, ontogenetic development and larval growth were near identical between triploids and diploids. However, under demanding conditions (i.e. the maximum swimming velocity during an escape response), triploid larvae performed poorer than their diploid counterparts, especially after repeated stimuli to induce swimming. This result is consistent with the idea that larger cells have less capacity to generate energy, which becomes manifest during repeated physical exertion resulting in increased fatigue. Triploidy induction in zebrafish appears a valid method to increase specifically cell size and this provides a model system to test for consequences of cell size adaptation for the energy budget and swimming performance of this ectothermic vertebrate.
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