Abstract
The notion that cultured fishes develop a morphology that differs from their wild conspecifics has become nearly axiomatic. A commonly supervened corollary is that exposure to culture causes predictable and consistent morphological changes that together form a common “cultured phenotype”. While this is often asserted, it has not been formally tested. We conducted a systematic review of the literature based on PRISMA best practice protocols and identified 65 papers, composed of 106 studies that compared the morphology of 39 species of cultured fish to their wild conspecifics. From this we conducted a meta-analysis of quantitative, and vote-counting analysis of qualitative differences in 16 external morphological features and condition factor. Our results confirm that aspects of a general “cultured phenotype” exist. The meta-analysis analysis revealed that cultured fish had shorter heads, upper jaws and fins, and the vote-counting analysis was suggestive of this as well. The vote-counting analysis showed that across all studies cultured fish had greater body depth and condition factor than wild fish, but this was not supported by the meta-analysis. In addition to matching those required to develop the “cultured phenotype”, the detected morphological changes are consistent with experimentally observed plastic responses to conditions typical of the cultured environment. This is discussed, as is how the observed changes may be influenced or reinforced by intentional and unintentional selection.
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Acknowledgments
We thank P. Arechavala-Lopez, S. Blanchet, M. Jahunen, K. Tiffan and T. Vehanen for providing us with data. The Natural Sciences and Engineering Research Council of Canada funded this research through a strategic grant to I.A.F. and C.F.P, and the Research and Development Corporation of Newfoundland provided additional support to B.F.W.
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Table S1 Keywords used in the systematic review. These keywords, along with Boolean operators, were used in relevant combinations to search the electronic databases. (PDF 24 kb)
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Table S2 References screened during the systematic review, along with the results of four inclusion criteria. (PDF 207 kb)
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Table S3 Summary of meta-analysis of qualitative morphological differences between cultured and wild fish. Heading abbreviations are as follows: LH is life history, Up is upper, Low is lower, K is Fulton’s condition factor, Caud Ped is caudal peduncle, Pec is pectoral, Pel is pelvic, L is length, D is depth, W is width, H is height, and F is fin. Within the table, Unk indicates that the data was not provided, or was ambiguous in the study. Comparisons of wild-caught fish to cultured are denoted WF, while CG indicates the fish were compared in a common garden. Different is abbreviated Diff. and Immature, Imm. C > W denotes studies in which the expression of the trait in cultured fish is greater than in the wild to which they were compared, C < W indicates the opposite and C = W indicates a trait was measured, but no difference was detected. Blank spaces signify that a trait was not measured in a given study. Species abbreviations and their corresponding common, and binomial names are listed. Where more than one comparison was conducted in a study, this is noted and the populations or comparison are noted. (DOCX 165 kb)
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Table S4 Summary of the results of the vote-counting analysis. Results are presented for each category of each moderator. Morph. Feature is short for morphological feature. Diff. w/i stands for difference within category, and is the result of the test of the hypothesis that the proportion studies finding each of the three possible relative differences in morphological feature size between the cultured and wild fish (i.e. C < W = C > W = C = W) are equal for a given morphological feature and category of a moderator. Where significant differences were found, the results of all pairwise comparisons and adjusted p-values are given. Diff. b/w stands for difference between categories, and is the result of the test of the hypothesis that for each of the three possible relative differences in morphological feature size, the proportion of studies finding it did not differ between categories of a moderator. Where significant, and there are more than two categories of a moderator, the results of all pairwise comparisons, and adjusted p-values reported. NS indicates a given test was not significant, and Chi squared and p-values are given for each test. Lab is short for laboratory, CG is common garden, and WF indicates studies in which the cultured fish were compared to wild-caught fish. There were a number of cases in which the sample size was not sufficient for accurate statistical analysis, and these are marked with an asterisk (*). In these cases it was impossible to test for Diff. b/w and this is left blank. (DOCX 219 kb)
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Table S5 Congruence of the results of the meta-analysis and the vote-counting analysis. For the meta-analysis the difference in size is what was indicated from the mean effect size as generated by the mixed-mixed effect model. For the vote-counting analysis, the relative difference found in the largest proportion of studies is used. Where the proportions of two possible differences did not differ, both are given, and where all three did not differ, this is denoted Aprx. Equal, or all proportions approximately equal. Significances are given for both the meta-analysis and the vote-counting results. Comp. is the comparison of the results of the vote-counting and meta-analysis. Where the results of the two analyses show the same relative difference in size for a morphological character between the cultured and wild fish the results are said to be congruent, where the two analyses show opposing differences, this is indicated as opposite, and where the they do not match, this is indicated as incongruent. A summary of the number of morphological features that were found to be congruent, incongruent or opposite are given for each moderator level. (DOCX 153 kb)
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Fig. S1 Forest-plots for each morphological feature examined. The points are the effect size for each study, and the error bars represent the 95% confidence interval around it. The size of the point is reflective of the weighting given to it by the linear-mixed effects function, and a unique colour is given to each genus. The morphological features are as described in Fig. 1/Table 1, and the species examined can be found in Table S3. S1a, Head depth; S1b, Head length; S1c, Eye size; S1d, Upper jaw length; S1e, Lower jaw length; S1f, Body depth; S1g, Condition factor; S1h, Caudle peduncle depth; S1i, Caudle peduncle length; S1j, Pectoral fin length; S1k, Pelvic fin length; S1l, Dorsal fin length; S1m, Dorsal fin width; S1n, Anal fin length; S1o, Anal fin width; S1p, Caudle fin length. (DOCX 3593 kb)
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Wringe, B.F., Purchase, C.F. & Fleming, I.A. In search of a “cultured fish phenotype”: a systematic review, meta-analysis and vote-counting analysis. Rev Fish Biol Fisheries 26, 351–373 (2016). https://doi.org/10.1007/s11160-016-9431-4
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DOI: https://doi.org/10.1007/s11160-016-9431-4