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BMC Evol Biol. To find out if the difference in mating frequency between N2 and HW hermaphrodites might represent an active process, we first manipulated the arena size of the assay. Here, we can suggest only three possible reasons. Further, the observation that females of C. remanei, a closely related obligate outcrossing species, are much more attractive to heterospecific C. elegans males than are conspecific C. elegans hermaphrodites (Chasnov et al. The numbers of attracted males (C. elegans or C. remanei) are shown; N is the number of independent trials; P value indicates the significance of the result. Furthermore, as shown in Table 2C, when the C. remanei female was killed by a hot worm pick the carcass did not elicit the same attraction, suggesting that the attractant was emitted only by live females. To characterize underlying neural mechanisms, we show that N2 hermaphrodites deficient in mechanosensation or chemosensation (e.g., mec-3 and osm-6 mutants) exhibit high mating frequency, implicating hermaphrodite perception of males as a requirement for low mating frequency. We find that males use a combination of volatile sex pheromones (VSPs), ascaroside pheromones, surface-bound chemical cues, and other signals to robustly assess a variety of features of potential mates. Abbreviated wiring diagram of sexually dimorphic neurons, Figure 2:. See this image and copyright information in PMC. Major anatomical features of a hermaphrodite (A) and male (B) viewed laterally. This addition completely realized the vision of C. elegans pioneer Sydney Brenner: a model organism with an entirely mapped nervous system. As a test of this hypothesis, we conducted a competition experiment using a wild-type N2 strain to compete against a mutant him-5 strain. By direct observation, successful copulation by males is very infrequent. Collet J, Spike C A, Lundquist E A, Shaw J E, Herman R K. Davis M W, Hammarlund M, Harrach T, Olsen S, Jorgensen E M. Kiontke K, Gavin N P, Raynes Y, Roehrig C, Piano F et al. Tracking N- and C-termini of C. elegans polycystin-1 reveals their distinct targeting requirements and functions in cilia and extracellular vesicles. Competition experiment between him-5 and wild type: The population competition experiment on C. elegans was performed in triplicate with the starting culture composed of 100 him-5 mutant hermaphrodites on a 100-mm culture plate. Reproductive evolution: symptom of a selfing syndrome. The role of hermaphrodites in the experimental evolution of increased outcrossing rates in Caenorhabditis elegans. 2007) suggests that early C. elegans hermaphrodites may have been more attractive or willing to mate with males than they are today. elegans is a non-hazardous, non-infectious, non-pathogenic, non-parasitic organism. Our parental strain, N2, had an average brood size of 195 26 (134266, N = 40), and AB1 had an average of 182 27 (148251, N = 50). (A) Hermaphrodite mating frequency of 158 recombinant inbred lines generated from a cross between N2 and HW (see Supporting Information, File S2). Recombinant inbred lines (RILs) were generated by intercrossing N2 and CB4856 (HW) and then selecting single hermaphrodite self-progeny for 710 generations from several hundred independent F2-derived lines, resulting in a panel of largely homozygous strains with mixed parental contribution from N2 and HW. WebC. The inability of uncoordinated hermaphrodites to migrate actively within the mating area enhances mating success 40- to 50-fold, and an increased density of hermaphrodites enhances mating success by 2-fold. The number of progeny produced by each individual was averaged over the total test worms to reflect the fecundity of the parental worms of specific genotype. Furthermore, any males present in the C. elegans population are under selection pressure to successfully mate with hermaphrodites, and we present theoretical results showing that the amount of outcrossing observed experimentally is sufficient to support male-specific genes against degeneration by deleterious mutations. Moreover, when tested in a separate experiment on very large plates (area = 175 cm2), HW hermaphrodites still exhibited relatively high mating frequency (63%, N = 24 pairs) compared to N2 (9%, N = 22 pairs) (P < 0.001), showing that HW hermaphrodites are highly prone to mating even when the encounter rate is likely to be low. Thus, as an assay for mating frequency, we placed one virgin L4 hermaphrodite and one virgin L4 male on standard laboratory plates (6-cm diameter with E. coli as food) and scored the sex of the progeny after 2 d of potential interaction (see File S2). Contrary to this prediction, we observed that HW had a largely similar tempo of self-reproduction compared to N2 (Figure 2B), and there was no obvious correlation with the appearance of self-sperm depletion and mating frequency (e.g., compare N2vs.HW during day 0 and day 1 treatments in Figure 2, A and B). The dioecious species had an average mating time of 41.6 33.2 min, and copulation was observed in 100% (40/40) of the matings. The evolution from females to hermaphrodites results in a sexual conflict over mating in androdioecious nematode worms and clam shrimp. 2009). First, we measured the correlation of natural sperm depletion and mating frequency in N2 and HW hermaphrodites. sharing sensitive information, make sure youre on a federal Ascarosides also signal nutritional information, allowing males to choose well-fed over starved mates, while both ascarosides and surface-associated cues cause males to prefer virgin over previously mated hermaphrodites. Curr Biol. Because the nematode C. elegans reproduces mostly by self-fertilization, little is known about its mate-choice behaviors. In fact, the ability of C. elegans to both self-fertilize and outcross has rendered it a powerful genetic model for developmental studies, and well-established laboratory procedures as well as readily available mutants facilitate experimentation. 2005), unlike unc-13 and unc-31 mutants described, it is unlikely that the high mating frequency of osm-6 mutants results from locomotion defects. To confirm this, we rescued the tax-4(ks28) mutant with a N2 wild-type tax-4 transgene (Figure S2). This NIL B exhibited low mating frequency, similar to N2 (Figure 3D), suggesting that the right half of the fragment is not sufficient to augment mating. Further, we found that the N2-derived npr-1(ad609) mutantwhich displays aggregation behavior similar to HWhad wild-type N2 hermaphrodite mating frequency (Figure S4). We begin with some words of caution. Here, we use a choice-based social-interaction assay to explore the ability of C. elegans males to make and optimize mate choices. We observed that both unc-13 and unc-31 mutant hermaphrodites exhibited high mating frequency, similar to HW (Figure 1D). 2006); thus, we used HW males as the tester strain in subsequent experiments (unless otherwise noted) to dissect hermaphrodite mating frequency. Our result is for the frequency u of spontaneous male births by C. elegans hermaphrodites. We measured u by counting 5695 males out of 17,488 offspring from virgin hermaphrodites, resulting in u = 0.3257 0.0035. We also show that the ability to self-reproduce negatively regulates hermaphrodite mating. As soon as hermaphrodite attractiveness became disadvantageous, any mutation that eliminated sex pheromone expression would have rapidly swept through the hermaphrodite population. Males can successfully mate with a hermaphrodite for 6 days after their last C. remanei mates more efficiently than C. elegans: In contrast to a hermaphroditic species, a dioecious species depends on successful male-female matings. Together, our findings lead to an integrated view of the signaling and behavioral mechanisms by which males use diverse sensory cues to assess multiple features of potential mates and optimize mate choice. The fractions of hermaphrodite and male zygotes from selfed and male-fertilized eggs are presented in Table 1. Hermaphrodites self-reproduce through internal self-fertilization with their own sperm and can outcross with males. If so, then why are there still males present in the C. elegans species? A mutation that results in a 50% increase in hermaphrodite sperm production is outcompeted by wild-type worms. We thank the Caenorhabditis Genetics Centerfunded by the National Institutes of Healthand the Bargmann Laboratory for strains; Zhunan Chen for custom-stratified permutation test software; Jesse Weber and Andres Bendesky for discussions about QTL mapping; Steve Buschbach, Colleen Carlston, Flavia Chen, Petra Kubikova, Nicholas Keone Lee, Chris Meehan, Manuel Ramos, Mary Stevens, Vivian Yeong, and Jingyi Yu for technical assistance; and members of the Zhang Laboratory for constructive discussions. A further simplifying approximation considers all the male-only expressed genes to be tightly linked. WebC. Intriguingly, we found that a more subtle mutation in tax-2, p694which causes loss of TAX-2 expression (and associated chemosensory responses) in just four pairs of amphid neurons because of a deletion in the tax-2 promoter, first exon, and first intron (Coburn and Bargmann 1996)exhibited high mating frequency (Figure 1H), a phenotype opposite of the strong loss-of-function tax-2 and tax-4 mutants described. elegans is an unsegmented pseudocoelomate and lacks respiratory or circulatory systems. (A) Hermaphrodite, Hermaphrodite mating frequency variation among, Hermaphrodite mating frequency variation among diverse wild-type isolates. The phylum nematoda provides us with two closely related species that may be relevant to the problem of the stability of the dioecious mating system: Caenorhabditis remanei and C. elegans. Genetic variants of C. elegans used in this study include AB1 (Australia). 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