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. 2021 Jul 24;38(11):4977–4986. doi: 10.1093/molbev/msab221

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© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.

This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial License (https://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact journals.permissions@oup.com

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Fig. 4 — Functional divergence of the D1 chromocenter forming protein from D. melanogaster and D. simulans. (A) D1 mutant (D1^LL03310) testes expressing HA-tagged UASt-D1^mel and UASt-D1^sim under the control of nos-gal4 were stained for Vasa (gray). Scale bars are 25 µm. (B) Quantification of HA+ germ cells in testes from 0- to 4-day-old and 12- to 16-day-old males of the following genotypes, nos>D1^mel; D1^LL03310 (n = 29, 0–4 days) and (n = 31, 12–16 days) and nos>D1^sim; D1^LL03310 (n = 31, 0–4 days) and (n = 32, 12–16 days). n indicates the number of testes analyzed, ** represents P = 0.006 (0–4 days) and P = 0.001 (12–16 days) from a Student’s t-test. (C) IF against Prod (magenta), HA (green), and Lamin (blue) in spermatogonial cells of the indicated genotypes. (D) Box-and-whisker plot of HA foci/cell in D1 mutant (D1^LL03310) spermatogonia expressing D1^mel-HA (n = 57) or D1^sim-HA (n = 60). n indicates the number of germ cells analyzed, **** represents P < 0.0001 from Student’s t-test and crosshairs mark the mean. (E) Box-and-whisker plot of Prod foci/cell in D1 mutant (D1^LL03310) spermatogonia expressing D1^mel-HA (n = 57) or D1^sim-HA (n = 60). n indicates the number of germ cells analyzed, **** represents P < 0.0001 from Student’s t-test and crosshairs mark the mean. (F) FISH against the (GAACAGAACATGTTC)_n satellite (magenta) in spermatogonial cells from the indicated 0- to 3-day-old males and costained with DAPI (blue) and Lamin (green). (G) Box-and-whisker plot of large autosomal (GAACAGAACATGTTC)_n foci per spermatogonial cell from D. sim C167.4 (n = 39) and D. sim C167.4×D. mau w⁺ (n = 31). n indicates the number of germ cells analyzed, ** represents P = 0.0067 from a Student’s t-test and crosshairs mark the mean. (H) IF against Lamin (green) in spermatogonial cells from the indicated 0- to 3-day-old males and costained with DAPI (magenta) and phalloidin (blue). Arrows point to micronuclei. (I) Quantification of testes containing at least one cell with a micronucleus from 0- to 4-day-old D. sim C167.4 pure species males (n = 18) and 0- to 4-day-old D. sim C167.4×D. mau hybrid males (n = 26). The percentage of micronuclei-containing testes is indicated above the respective columns. n indicates the number of testes analyzed, ** represents P = 0.0027 from Fisher’s exact test. All scale bars (except panel A) are 5 µm and yellow dashed lines demarcate nuclear boundary. (J) A speculative model of how hybrid incompatibility factors may inhibit chromocenter formation by species-specific satellite DNA-binding proteins in cells of affected hybrid tissues.