38:
679:, the egg is fertilized. Social bees, wasps, and ants can modify sex ratios within colonies which maximizes relatedness among members and generates a workforce appropriate to surrounding conditions. In other solitary hymenopterans, the females lay unfertilized male eggs on poorer food sources while laying the fertilized female eggs on better food sources, possibly because the fitness of females will be more adversely affected by shortages in their early life. Sex ratio manipulation is also practiced by haplodiploid
692:
2616:
804:, a non-CSD method of sex determination has been documented. The most recent accepted model for this non-CSD system is called Maternal Effect Genomic Imprinting Sex Determination (MEGISD). This model involves a masculinizing/virilizing maternal effect gene that “imprints upon” the cytoplasmic component of oocytes, and an “unimprinted” paternal contribution (in female offspring) that provides a counter effect to
88:
2189:
776:
is possibly a strategy to ensure the proper working of the hive. However, since queens usually mate with a dozen drones or more, not all workers are full sisters. Due to the separate storage of drone sperm, a specific batch of brood may be more closely related than a specific batch of brood laid at a
721:
cells that contain their entire genome, so the sperm are all genetically identical except for mutations. The male bees' genetic makeup is therefore entirely derived from the mother, while the genetic makeup of the female worker bees is half derived from the mother, and half from the father. Thus, if
808:
and allows for female development to occur. Since all diploid eggs become female (due to the factor originating in the male genetic contribution that prevents masculinization), this differs from CSD in that under CSD, diploid eggs can become males if they are homozygous or hemizygous.
768:
due to indirect selection. Since the worker is more related to the queen's daughters (her sisters) than to her own offspring, helping the queen's offspring to survive helps the spread of the same genes that the worker possesses more efficiently than direct reproduction.
1595:
Van Alphen, J. J. M.; Thunnissen, I. (1982). "Host
Selection and Sex Allocation by Pachycrepoideus Vindemiae Rondani (Pteromalidae) as a Facultative Hyperparasitoid of Asobara Tabida Nees (Braconidae; Alysiinae) and Leptopilina Heterotoma (Cynipoidea; Eucoilidae)".
743:
There are rare instances of diploid drone larvae. This phenomenon usually arises when there is more than two generations of brother-sister mating. Sex determination in honey bees is initially due to a single locus, called the complementary sex determiner
760:
gene, they will develop into males. The instances where the individual is homozygous at this gene are the instances of diploid males. Diploid males do not survive to adulthood, as the nurse worker bees will cannibalize the diploid males upon hatching.
600:
will be removed from the population rapidly because they will automatically be expressed in the males (dominant lethal and deleterious alleles are removed from the population every time they arise, as they kill any individual they arise in).
612:. In an X0 sex-determination system, males and females receive an equal number of autosomes, but when it comes to sex chromosomes, females will receive two X chromosomes while males will receive only a single X chromosome.
893:, all her daughters share a complete set of chromosomes from that male. In Hymenoptera, the males generally produce enough sperm to last the female for her whole lifetime after a single mating event with that male.
1234:. However, colonies which have workers from multiple queens or queens which have mated multiple times will have worker-to-worker relatedness which is less than worker-to-daughter relatedness, such as in
579:. This means the workers in such monogamous single-queen colonies are significantly more closely related than in other sex determination systems where the relatedness of siblings is usually no more than
2443:
644:
offspring develop from fertilized eggs, and are normally female, while haploid offspring develop into males from unfertilized eggs. Diploid males would be infertile, as their cells would not undergo
561:
The haplodiploid sex-determination system has a number of peculiarities. For example, a male has no father and cannot have sons, but he has a grandfather and can have grandsons. Additionally, if a
787:. This means that sisters are almost always more related to one another than they would be to their own offspring, thus eliminating the conflict of variable relatedness present in honeybees.
896:
Relatedness coefficients in haplodiploid organisms are as follows, assuming that a female has only mated once. These ratios apply, for example, throughout a bee hive, unless some
2436:
1845:
1230:
Under this assumption that mothers only mate once, sisters are more strongly related to each other than to their own daughters. This fact has been used to explain the
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554:
develops as a female, and an unfertilized egg develops as a male. This means that the males have half the number of chromosomes that a female has, and are
1667:
842:
behavior in hymenopterans. A eusocial worker helping her mother birth more sisters propagates more of her own genes than had she reproduced herself.
1858:
Zou, Yuan et al. “A chimeric gene paternally instructs female sex determination in the haplodiploid wasp
Nasonia.” Science 370 (2020): 1115 - 1118.
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149:
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theory of how eusociality evolved. Whether haplodiploidy did in fact pave the way for the evolution of eusociality is still a matter of debate.
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Several models have been proposed for the genetic mechanisms of haplodiploid sex-determination. The model most commonly referred to is the
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1687:
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While workers can lay unfertilized eggs that become their sons, haplodiploid sex-determination system increases the individual's
403:
238:
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1560:
Chow, A.; MacKauer, M. (1996). "Sequential allocation of offspring sexes in the hyperparasitoid wasp, Dendrocerus carpenteri".
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produce offspring, which will all be males from unfertilised eggs: in that case, average relatedness will be lower than shown.
675:. The mated female controls the release of stored sperm from within the organ: If she releases sperm as an egg passes down her
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208:
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Baer, B.; P. Schmid-Hempel (2001). "Unexpected consequences of polyandry for parasitism and fitness in the bumblebee,
1726:
Cannibalism of
Diploid Drone Larvae in the Honey Bee (Apis mellifera) is Released by Odd Pattern of Circular Substance
683:, who lay more male eggs when the chances for males to disperse and mate with females in different sites are greater.
605:
144:
139:
134:
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Hughes, W.O.H.; et al. (2008). "Ancestral monogamy shows kin selection is key to the evolution of eusociality".
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for her daughters, but the haploid father's genome is inherited by his daughters "as is". It is also possible for a
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are unusually closely related to their full sisters (same father) because of their haplodiploid inheritance system.
341:
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717:, their mother. The diploid queen has 32 chromosomes and the haploid drones have 16 chromosomes. Drones produce
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1625:"Female ambrosia beetles adjust their offspring sex ratio according to outbreeding opportunities for their sons"
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relatedness coefficient amongst full haplodiploid sisters is responsible for the frequency of evolution of
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1989:"Single-locus complementary sex determination absent in Heterospilus prosopidis (Hymenoptera: Braconidae)"
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309:
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107:
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Ratnieks, Francis (1988). "Reproductive harmony via mutual policing by workers in eusocial hymenoptera".
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1743:"The Effect of Sex-Allocation Biasing on the Evolution of Worker Policing in Hymenopteran Societies"
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Batches of worker bees are short lived and are constantly being replaced by the next batch, so this
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to form sperm. Therefore, the sperm would be diploid, which means that their offspring would be
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652:. Since hymenopteran mother and sons share the same genes, they may be especially sensitive to
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608:. In haplodiploidy, males receive one half of the chromosomes that females receive, including
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2010:
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748:) gene. In developing bees, if the conditions are that the individual is heterozygous for the
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Weinstock, George M.; Robinson, Gene E., & the
Honeybee Genome Sequencing Consortium
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a queen bee mates with only one drone, any two of her daughters will share, on average,
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1970:
1532:
Single locus complementary sex determination in
Hymenoptera: an "unintelligent" design?
171:
1940:"Unusually high recombination rate detected in the sex locus region of the honey bee (
1939:
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gene, they will develop into females. If the conditions are so that the individual is
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Edward O. Wilson (2005). "Kin
Selection as the Key to Altruism: Its Rise and Fall".
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Shared gene proportions in haplo-diploid sex-determination system relationships
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later date. However, many other species of bees, including bumblebees, such as
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Biological system where sex is determined by the number of sets of chromosomes
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1367:
White, Michael J.D. (1984). "Chromosomal mechanisms in animal reproduction".
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Insights into social insects from the genome of the honeybee Apis mellifera
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present in a population, hence increasing the occurrence of diploid males.
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565:-insect colony has only one queen, and she has only mated once, then the
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889:). So, all female offspring inherit the male's chromosomes 100% intact.
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Narrow roads of gene land : the collected papers of W.D. Hamilton
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Haplodiploidy determines the sex in all members of the insect orders
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Santomauro, Giulia; Oldham, Neil J.; Boland, Wilhelm; Engels Wolf;
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36:
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van
Wilgenburg, Ellen; Driessen, Gerard & Beukeboom, Leo W.
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an individual receives. An offspring formed from the union of a
512:
2425:
2062:
1896:
Kevin R. Foster; Tom
Wenseleers; Francis L.W. Ratnieks (2006).
1830:
Davies, Nicholas B., John R. Krebs and Stuart A. West. (2012).
869:, because in each chromosome of a pair there will be different
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Another feature of the haplodiploidy system is that recessive
508:
504:
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1809:
10.1554/0014-3820(2001)055[1639:ucopfp]2.0.co;2
542:
In this system, sex is determined by the number of sets of
828:). The haplodiploidy hypothesis proposes that the unusual
519:('thrips'). The system also occurs sporadically in some
569:
between workers (diploid females) in a hive or nest is
484:
in which males develop from unfertilized eggs and are
1665:
Kin
Selection and Haplodiploidy in Social Hymenoptera
1298:
King, R.C.; Stansfield, W.D.; Mulligan, P.K. (2006).
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to lay an unfertilised egg, which is always a male.
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488:, and females develop from fertilized eggs and are
1741:Foster, Kevin R.; Ratnieks, Francis L. W. (2001).
1685:Drone Larvae from Fertilized Eggs of the Honey Bee
1306:(7th ed.). Oxford University Press. p.
656:: Inbreeding reduces the number of different sex
891:As long as a female has mated with only one male
640:individuals develop into males. In other words,
624:. According to this model, if an individual is
34:(a haploidized diploid or diploidized haploid).
1232:evolution of eusociality in many hymenopterans
849:, the father has two sets of chromosomes, and
732:of their genes. The diploid queen's genome is
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8:
1844:: CS1 maint: multiple names: authors list (
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2422:
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1875:. Oxford New York: W.H. Freeman/Spektrum.
667:, each fertile hymenopteran female stores
604:Haplodiploidy is not the same thing as an
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2004:
1969:
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2483:Development of the reproductive system
1898:"Kin selection is the key to altruism"
1837:
1715:, volume "'443'" (2006), pages 931-949
1543:Mahowald, Michael; von Wettberg, Eric
1410:. Cambridge University Press. p.
1339:. Cambridge University Press. p.
713:(males) are entirely derived from the
1832:An Introduction to Behavioral Ecology
820:is used to calculate the strength of
632:, it develops into a female, whereas
7:
1834:. Wiley-Blackwell. pp. 371–375.
1545:Sex determination in the Hymenoptera
589:. It is this point which drives the
492:. Haplodiploidy is sometimes called
1731:, volume "'43'" (2004), pages 69-74
813:Relatedness ratios in haplodiploidy
1938:Beye, Martin; et al. (1999).
1683:Woyka, J.; Pszczelnictwa, Zaklad;
25:
1902:Trends in Ecology & Evolution
1402:Grimaldi, D.; Engel M.S. (2005).
1331:Grimaldi, D.; Engel M.S. (2005).
881:(except for a small number where
877:. But when the father is haploid
2615:
2614:
2187:
1642:10.1111/j.1420-9101.2003.00687.x
239:Evolution of sexual reproduction
86:
2391:Sociobiology: The New Synthesis
1629:Journal of Evolutionary Biology
1623:Peer, K.; Taborsky, M. (2004).
687:Sex determination in honey bees
1729:Journal of Apiculture Research
1695:Journal of Apiculture Research
1598:Netherlands Journal of Zoology
1534:Frontiers in Zoology 2006, 3:1
671:in an internal sac called the
598:lethal and deleterious alleles
332:Sexual reproduction in animals
1:
2597:Disorders of sex development
1987:Wu, Z.; et al. (2005).
1406:The evolution of the insects
1335:The evolution of the insects
879:all the sperms are identical
867:the sperms are not identical
209:Simultaneous hermaphroditism
2157:Sexual selection in insects
1962:10.1093/genetics/153.4.1701
1173:Niece (brother's daughter)
606:X0 sex-determination system
2672:
2309:Austroplatypus incompertus
2137:Identity in social insects
1914:10.1016/j.tree.2005.11.020
1146:Niece (sister's daughter)
622:complementary allele model
342:Penile-vaginal intercourse
204:Sequential hermaphroditism
29:
2641:Sex-determination systems
2610:
2185:
1381:10.1080/11250008409439455
194:Testis-determining factor
2516:Sex-determination system
2104:Evolution of eusociality
1871:Hamilton, W. D. (1996).
1302:A dictionary of genetics
885:have taken place in the
857:of each pair during the
853:takes place between the
482:sex-determination system
187:Sex chromosome anomalies
125:Sex-determination system
47:sex-determination system
30:Not to be confused with
2367:Charles Duncan Michener
1750:The American Naturalist
1610:10.1163/002829683X00228
1459:10.1126/science.1156108
2564:Sex determining gene:
2471:Sexual differentiation
2006:10.1038/sj.hdy.6800720
1574:10.1006/anbe.1996.0090
1369:Bollettino di Zoologia
1265:Sexual differentiation
1200:Nephew (sister's son)
798:In wasps of the genus
756:or homozygous for the
702:
310:Internal fertilization
305:External fertilization
108:Sexual differentiation
62:
2536:Temperature-dependent
2372:The Bees of the World
1697:, (1963), pages 19-24
1502:10.1353/sor.2005.0012
791:Sex determination in
694:
150:Temperature-dependent
40:
2500:Paramesonephric duct
1237:Melipona scutellaris
57:females. System for
2407:Journey to the Ants
2024:American Naturalist
1451:2008Sci...320.1213H
1445:(5880): 1213–1216.
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861:which produces the
847:sexual reproduction
327:Fungal reproduction
230:Sexual reproduction
1690:2014-10-22 at the
1670:2011-07-16 at the
1548:Swarthmore College
1260:Pseudo-arrhenotoky
1250:Green-beard effect
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354:Human reproduction
337:Sexual intercourse
322:Plant reproduction
216:Intersex (biology)
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2333:Bees in mythology
1882:978-0-7167-4530-3
1793:Bombus terrestris
1421:978-0-521-82149-0
1350:978-0-521-82149-0
1317:978-0-19-530761-0
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780:Bombus terrestris
738:laying worker bee
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359:Lordosis behavior
162:Heterogametic sex
103:Sexual dimorphism
16:(Redirected from
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2347:Pioneers, works
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2162:Social conflict
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2127:Group selection
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2623:
2622:
2611:
2608:
2607:
2605:
2604:
2599:
2594:
2589:
2583:
2581:
2577:
2576:
2574:
2573:
2562:
2561:
2560:
2555:
2548:Sex chromosome
2545:
2544:
2543:
2538:
2533:
2528:
2523:
2512:
2510:
2506:
2505:
2503:
2502:
2497:
2492:
2491:
2490:
2480:
2479:
2478:
2467:
2465:
2461:
2460:
2451:
2449:
2448:
2441:
2434:
2426:
2417:
2416:
2414:
2413:
2412:
2411:
2403:
2395:
2382:
2377:
2376:
2375:
2364:
2363:
2362:
2350:
2348:
2344:
2343:
2341:
2340:
2335:
2329:
2327:
2323:
2322:
2320:
2319:
2314:
2313:
2312:
2300:
2299:
2298:
2288:
2287:
2286:
2274:
2273:
2272:
2260:
2259:
2258:
2253:
2251:Dwarf mongoose
2248:
2238:
2237:
2236:
2231:
2226:
2221:
2216:
2211:
2200:
2198:
2194:
2193:
2186:
2184:
2182:
2181:
2180:
2179:
2174:
2169:
2164:
2159:
2154:
2149:
2144:
2139:
2134:
2129:
2124:
2119:
2111:
2106:
2100:
2098:
2094:
2093:
2088:
2086:
2085:
2078:
2071:
2063:
2057:
2056:
2036:10.1086/284846
2030:(2): 217–236.
2019:
1999:(3): 228–234.
1984:
1942:Apis mellifera
1933:
1930:
1928:
1927:
1888:
1881:
1860:
1851:
1822:
1783:
1762:10.1086/323588
1756:(6): 615–623.
1733:
1717:
1699:
1676:
1656:
1635:(2): 257–264.
1615:
1604:(4): 497–514.
1587:
1568:(4): 859–870.
1552:
1536:
1523:
1496:(1): 159–166.
1480:
1427:
1420:
1394:
1356:
1349:
1323:
1316:
1289:
1287:
1284:
1283:
1282:
1277:
1272:
1267:
1262:
1257:
1252:
1245:
1242:
1228:
1227:
1224:
1223:
1212:
1201:
1197:
1196:
1185:
1174:
1170:
1169:
1158:
1147:
1143:
1142:
1139:
1128:
1124:
1123:
1120:
1109:
1108:Paternal Aunt
1105:
1104:
1093:
1082:
1078:
1077:
1066:
1055:
1054:Maternal Aunt
1051:
1050:
1039:
1028:
1024:
1023:
1012:
1001:
997:
996:
993:
982:
978:
977:
974:
963:
959:
958:
955:
944:
940:
939:
936:
925:
921:
920:
917:
914:
898:laying workers
883:gene mutations
814:
811:
795:
789:
688:
685:
628:for a certain
617:
614:
473:
472:
470:
469:
462:
455:
447:
444:
443:
442:
441:
436:
431:
423:
422:
416:
415:
414:
413:
412:
411:
406:
401:
391:
386:
378:
377:
371:
370:
369:
368:
367:
366:
361:
356:
351:
346:
345:
344:
329:
324:
319:
314:
313:
312:
307:
297:
296:
295:
290:
280:
279:
278:
273:
263:
258:
253:
252:
251:
246:
233:
232:
226:
225:
224:
223:
218:
213:
212:
211:
206:
196:
191:
190:
189:
184:
179:
172:Sex chromosome
169:
164:
159:
158:
157:
152:
147:
142:
137:
132:
122:
121:
120:
115:
105:
97:
96:
92:
91:
83:
82:
74:
73:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2668:
2657:
2654:
2652:
2649:
2647:
2644:
2642:
2639:
2638:
2636:
2621:
2613:
2612:
2609:
2603:
2600:
2598:
2595:
2593:
2590:
2588:
2587:Hermaphrodite
2585:
2584:
2582:
2578:
2571:
2567:
2563:
2559:
2556:
2554:
2551:
2550:
2549:
2546:
2542:
2541:Haplodiploidy
2539:
2537:
2534:
2532:
2529:
2527:
2524:
2522:
2519:
2518:
2517:
2514:
2513:
2511:
2509:Genetic basis
2507:
2501:
2498:
2496:
2493:
2489:
2486:
2485:
2484:
2481:
2477:
2474:
2473:
2472:
2469:
2468:
2466:
2462:
2458:
2454:
2447:
2442:
2440:
2435:
2433:
2428:
2427:
2424:
2409:
2408:
2404:
2401:
2400:
2396:
2393:
2392:
2388:
2387:
2386:
2383:
2381:
2378:
2373:
2370:
2369:
2368:
2365:
2360:
2357:
2356:
2355:
2352:
2351:
2349:
2345:
2339:
2336:
2334:
2331:
2330:
2328:
2324:
2318:
2315:
2311:
2310:
2306:
2305:
2304:
2301:
2297:
2294:
2293:
2292:
2289:
2285:
2284:
2280:
2279:
2278:
2275:
2271:
2270:
2266:
2265:
2264:
2261:
2257:
2254:
2252:
2249:
2247:
2244:
2243:
2242:
2239:
2235:
2232:
2230:
2227:
2225:
2222:
2220:
2217:
2215:
2212:
2210:
2207:
2206:
2205:
2202:
2201:
2199:
2195:
2190:
2178:
2175:
2173:
2170:
2168:
2165:
2163:
2160:
2158:
2155:
2153:
2150:
2148:
2147:Kin selection
2145:
2143:
2140:
2138:
2135:
2133:
2132:Haplodiploidy
2130:
2128:
2125:
2123:
2120:
2118:
2115:
2114:
2112:
2110:
2107:
2105:
2102:
2101:
2099:
2095:
2091:
2084:
2079:
2077:
2072:
2070:
2065:
2064:
2061:
2053:
2049:
2045:
2041:
2037:
2033:
2029:
2025:
2020:
2016:
2012:
2007:
2002:
1998:
1994:
1990:
1985:
1981:
1977:
1972:
1967:
1963:
1959:
1955:
1951:
1950:
1945:
1943:
1936:
1935:
1931:
1923:
1919:
1915:
1911:
1907:
1903:
1899:
1892:
1889:
1884:
1878:
1874:
1867:
1865:
1861:
1855:
1852:
1847:
1841:
1833:
1826:
1823:
1818:
1814:
1810:
1806:
1802:
1798:
1794:
1787:
1784:
1779:
1775:
1771:
1767:
1763:
1759:
1755:
1751:
1744:
1737:
1734:
1730:
1727:
1721:
1718:
1714:
1713:
1709:
1703:
1700:
1696:
1693:
1689:
1686:
1680:
1677:
1673:
1669:
1666:
1660:
1657:
1652:
1648:
1643:
1638:
1634:
1630:
1626:
1619:
1616:
1611:
1607:
1603:
1599:
1591:
1588:
1583:
1579:
1575:
1571:
1567:
1563:
1556:
1553:
1549:
1546:
1540:
1537:
1533:
1527:
1524:
1519:
1515:
1511:
1507:
1503:
1499:
1495:
1491:
1484:
1481:
1476:
1472:
1468:
1464:
1460:
1456:
1452:
1448:
1444:
1440:
1439:
1431:
1428:
1423:
1417:
1413:
1408:
1407:
1398:
1395:
1390:
1386:
1382:
1378:
1375:(1–2): 1–23.
1374:
1370:
1363:
1361:
1357:
1352:
1346:
1342:
1337:
1336:
1327:
1324:
1319:
1313:
1309:
1304:
1303:
1294:
1291:
1285:
1281:
1278:
1276:
1273:
1271:
1268:
1266:
1263:
1261:
1258:
1256:
1253:
1251:
1248:
1247:
1243:
1241:
1239:
1238:
1233:
1213:
1202:
1199:
1198:
1186:
1175:
1172:
1171:
1159:
1148:
1145:
1144:
1140:
1129:
1126:
1125:
1121:
1110:
1107:
1106:
1094:
1083:
1080:
1079:
1067:
1056:
1053:
1052:
1040:
1029:
1026:
1025:
1013:
1002:
999:
998:
994:
983:
980:
979:
975:
964:
961:
960:
956:
945:
942:
941:
937:
926:
923:
922:
918:
915:
912:
911:
908:
903:
902:
901:
899:
894:
892:
888:
884:
880:
876:
872:
868:
865:. Therefore,
864:
860:
856:
852:
851:crossing over
848:
843:
841:
827:
823:
822:kin selection
819:
812:
810:
807:
803:
802:
794:
790:
788:
786:
782:
781:
775:
774:kin selection
770:
767:
762:
759:
755:
751:
747:
741:
739:
735:
720:
716:
712:
708:
700:
697:
693:
686:
684:
682:
678:
674:
670:
666:
661:
659:
655:
651:
647:
643:
639:
635:
631:
627:
623:
615:
613:
611:
607:
602:
599:
594:
592:
591:kin selection
568:
564:
559:
557:
553:
549:
545:
540:
538:
534:
530:
526:
522:
518:
514:
510:
506:
502:
497:
495:
491:
487:
483:
479:
478:Haplodiploidy
468:
463:
461:
456:
454:
449:
448:
446:
445:
440:
439:Hermaphrodite
437:
435:
432:
430:
427:
426:
425:
424:
421:
420:Sexual system
417:
410:
407:
405:
402:
400:
397:
396:
395:
392:
390:
387:
385:
382:
381:
380:
379:
376:
372:
365:
364:Pelvic thrust
362:
360:
357:
355:
352:
350:
347:
343:
340:
339:
338:
335:
334:
333:
330:
328:
325:
323:
320:
318:
315:
311:
308:
306:
303:
302:
301:
300:Fertilization
298:
294:
291:
289:
286:
285:
284:
281:
277:
274:
272:
269:
268:
267:
266:Gametogenesis
264:
262:
259:
257:
254:
250:
247:
245:
242:
241:
240:
237:
236:
235:
234:
231:
227:
222:
219:
217:
214:
210:
207:
205:
202:
201:
200:
199:Hermaphrodite
197:
195:
192:
188:
185:
183:
180:
178:
175:
174:
173:
170:
168:
165:
163:
160:
156:
155:Haplodiploidy
153:
151:
148:
146:
143:
141:
138:
136:
133:
131:
128:
127:
126:
123:
119:
116:
114:
111:
110:
109:
106:
104:
101:
100:
99:
98:
93:
89:
85:
84:
81:
80:
75:
71:
67:
66:
60:
56:
52:
48:
44:
39:
33:
19:
2602:Sex reversal
2558:Y chromosome
2553:X chromosome
2540:
2405:
2397:
2389:
2385:E. O. Wilson
2371:
2358:
2307:
2281:
2277:Thysanoptera
2267:
2131:
2109:Presociality
2027:
2023:
1996:
1992:
1953:
1947:
1941:
1932:Bibliography
1908:(2): 57–60.
1905:
1901:
1891:
1872:
1854:
1831:
1825:
1800:
1796:
1792:
1786:
1753:
1749:
1736:
1720:
1710:
1702:
1679:
1659:
1632:
1628:
1618:
1601:
1597:
1590:
1565:
1561:
1555:
1539:
1526:
1493:
1489:
1483:
1442:
1436:
1430:
1405:
1397:
1372:
1368:
1334:
1326:
1301:
1293:
1280:Y chromosome
1275:X chromosome
1235:
1229:
906:
895:
890:
878:
866:
844:
816:
806:virilization
799:
797:
778:
771:
763:
757:
749:
745:
742:
704:
662:
626:heterozygous
621:
619:
603:
595:
560:
541:
533:bark beetles
521:spider mites
517:Thysanoptera
498:
477:
476:
288:spermatozoon
182:Y chromosome
177:X chromosome
154:
118:Virilization
113:Feminization
77:
18:Haplodiploid
2651:Hymenoptera
2283:Kladothrips
2219:Crabronidae
2204:Hymenoptera
2117:Agriculture
2090:Eusociality
818:Relatedness
673:spermatheca
567:relatedness
544:chromosomes
501:Hymenoptera
494:arrhenotoky
434:Gonochorism
221:Mating type
43:Hymenoptera
2646:Beekeeping
2635:Categories
2338:Coalescent
2326:In culture
2303:Coleoptera
2269:Synalpheus
2263:Crustacean
2224:Halictidae
2152:Polyethism
1286:References
855:chromatids
845:In normal
785:monandrous
754:hemizygous
734:recombined
654:inbreeding
638:homozygous
634:hemizygous
616:Mechanisms
529:Coleoptera
349:Copulation
53:males and
2568:(mammal)
2296:Aphididae
2291:Hemiptera
2229:Honey bee
2167:Thelytoky
2122:Gamergate
1840:cite book
1797:Evolution
1518:142713581
1389:0373-4137
924:Daughter
887:germ line
707:honeybees
696:Honey bee
610:autosomes
525:Hemiptera
399:Mechanics
375:Sexuality
276:Oogenesis
256:Germ cell
244:Anisogamy
59:honey bee
49:involves
32:Dihaploid
2620:Category
2592:Intersex
2580:See also
2464:Overview
2399:The Ants
2317:Isoptera
2241:Mammalia
2234:Vespidae
2052:84742198
2015:16077738
1993:Heredity
1980:10581277
1949:Genetics
1922:16701471
1817:11580023
1778:46052954
1770:18707355
1688:Archived
1668:Archived
1651:15009259
1582:53192398
1510:40972006
1475:20388889
1467:18511689
1244:See also
1027:Brother
840:eusocial
650:triploid
563:eusocial
537:rotifers
409:Activity
70:a series
68:Part of
2572:(birds)
2256:Meerkat
2246:Blesmol
2044:2461867
1971:1460844
1447:Bibcode
1438:Science
1218:⁄
1207:⁄
1191:⁄
1180:⁄
1164:⁄
1153:⁄
1134:⁄
1115:⁄
1099:⁄
1088:⁄
1072:⁄
1061:⁄
1045:⁄
1034:⁄
1018:⁄
1007:⁄
1000:Sister
988:⁄
981:Father
969:⁄
962:Mother
950:⁄
931:⁄
916:Female
871:alleles
859:meiosis
833:⁄
801:Nasonia
766:fitness
727:⁄
699:workers
677:oviduct
658:alleles
646:meiosis
642:diploid
584:⁄
574:⁄
556:haploid
550:and an
535:), and
490:diploid
486:haploid
261:Meiosis
249:Isogamy
55:diploid
51:haploid
41:In the
2488:gonads
2476:humans
2214:Apidae
2197:Groups
2097:Topics
2050:
2042:
2013:
1978:
1968:
1920:
1879:
1815:
1776:
1768:
1712:Nature
1649:
1580:
1550:(1999)
1516:
1508:
1473:
1465:
1418:
1387:
1347:
1314:
1255:Ploidy
783:, are
711:drones
709:, the
665:mating
663:After
515:) and
511:, and
429:Dioecy
283:Gamete
61:shown.
45:, the
2570:DMRT1
2048:S2CID
2040:JSTOR
1774:S2CID
1746:(PDF)
1578:S2CID
1514:S2CID
1506:JSTOR
1471:S2CID
919:Male
863:sperm
824:(via
719:sperm
715:queen
669:sperm
630:locus
548:sperm
513:wasps
480:is a
2455:and
2410:1994
2402:1990
2394:1975
2374:2000
2361:1927
2011:PMID
1976:PMID
1918:PMID
1877:ISBN
1846:link
1813:PMID
1766:PMID
1674:1997
1647:PMID
1463:PMID
1416:ISBN
1385:ISSN
1345:ISBN
1312:ISBN
943:Son
913:Sex
875:loci
636:and
509:ants
505:bees
293:ovum
2566:SRY
2209:Ant
2172:War
2032:doi
2028:132
2001:doi
1966:PMC
1958:doi
1954:153
1910:doi
1805:doi
1795:".
1758:doi
1754:158
1637:doi
1606:doi
1570:doi
1498:doi
1455:doi
1443:320
1412:465
1377:doi
1341:408
1308:194
758:csd
750:csd
746:csd
705:In
552:egg
79:Sex
2637::
2531:ZW
2526:X0
2521:XY
2046:.
2038:.
2026:.
2009:.
1997:95
1995:.
1991:.
1974:.
1964:.
1952:.
1946:.
1944:)"
1916:.
1906:21
1904:.
1900:.
1863:^
1842:}}
1838:{{
1811:.
1801:55
1799:.
1772:.
1764:.
1752:.
1748:.
1645:.
1633:17
1631:.
1627:.
1602:33
1600:.
1576:.
1566:51
1564:.
1512:.
1504:.
1494:72
1492:.
1469:.
1461:.
1453:.
1441:.
1414:.
1383:.
1373:51
1371:.
1359:^
1343:.
1310:.
1240:.
1141:—
1122:—
995:—
976:1
957:—
938:1
558:.
539:.
527:,
523:,
507:,
496:.
145:ZO
140:ZW
135:XO
130:XY
72:on
2445:e
2438:t
2431:v
2082:e
2075:t
2068:v
2054:.
2034::
2017:.
2003::
1982:.
1960::
1924:.
1912::
1885:.
1848:)
1819:.
1807::
1780:.
1760::
1653:.
1639::
1612:.
1608::
1584:.
1572::
1520:.
1500::
1477:.
1457::
1449::
1424:.
1391:.
1379::
1353:.
1320:.
1220:4
1216:1
1209:8
1205:3
1193:2
1189:1
1182:4
1178:1
1166:4
1162:1
1155:8
1151:3
1136:4
1132:1
1117:4
1113:1
1101:4
1097:1
1090:8
1086:1
1074:4
1070:3
1063:8
1059:3
1047:2
1043:1
1036:4
1032:1
1020:2
1016:1
1009:4
1005:3
990:2
986:1
971:2
967:1
952:2
948:1
933:2
929:1
835:4
831:3
744:(
729:4
725:3
586:2
582:1
576:4
572:3
531:(
503:(
466:e
459:t
452:v
20:)
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