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Multiple Paternity and Sporophytic Inbreeding Depression in a Dioicous Moss Species

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Multiple Paternity and Sporophytic Inbreeding Depression in a Dioicous Moss Species Heredity (2009) 103, 394–403 & 2009 Macmillan Publishers Limited All rights reserved 0018-067X/09 $32.00 www.nature.com/hdy ORIGINAL ARTICLE Multiple paternity and sporophytic inbreeding depression in a dioicous moss species P Szo¨ve´nyi, M Ricca and AJ Shaw Department of Biology, Duke University, Durham, NC, USA Multiple paternity (polyandry) frequently occurs in an average fixation coefficient and population level selfing flowering plants and animals and is assumed to have an rate of zero. In line with the prediction of sporophytic important function in the evolution of reproductive traits. inbreeding depression sporophyte size was significantly Polyandry in bryophytes may occur among multiple correlated with the level of heterozygosity. Furthermore, sporophytes of a female gametophyte; however, its occur- female gametophytes preferentially supported sporophytes rence and extent is unknown. In this study we investigate the with higher heterozygosity. These results indicate that occurrence and extent of multiple paternity, spatial genetic polyandry provides the opportunity for postfertilization selec- structure, and sporophytic inbreeding depression in tion in bryophytes having short fertilization distances natural populations of a dioicous bryophyte species, Sphag- and spatially structured populations facilitating inbreeding. num lescurii, using microsatellite markers. Multiple paternity Preferential maternal support of the more heterozygous is prevalent among sporophytes of a female gameto- sporophytes suggests active inbreeding avoidance that phyte and male genotypes exhibit significant skew in may have significant implications for mating system evolution paternity. Despite significant spatial genetic structure in the in bryophytes. population, suggesting frequent inbreeding, the number of Heredity (2009) 103, 394–403; doi:10.1038/hdy.2009.82; inbred and outbred sporophytes was balanced, resulting in published online 22 July 2009 Keywords: bryophytes; inbreeding; mating systems; reproductive skew; reproductive compensation; selective embryo abortion Introduction limited by pollen transport (Snow and Lewis, 1993; Campbell, 1998; Bernasconi, 2003; Knight et al., 2005). Male and female fecundity are often differently limited However, the effect of polyandry on female and male and this is thought to have led to a sexual conflict in fitness, gene flow, and the evolution of reproductive animals (Trivers, 1972). Female fecundity is expected to structures in flowering plants have started to receive be limited by egg production whereas male fecundity is attention only in the past few years (Bernasconi, 2003; usually limited by the number of mating events (Bate- Teixeira and Bernasconi, 2007; Teixeira et al., 2009). In man, 1948; Arnold, 1994). This has crystallized to the flowering plants, polyandry may increase overall female general idea that female animals might maximize their fitness of an individual either through pre- or post- fitness by mating with one male over a reproductive pollination selection. Mating with multiple mates may cycle. Nevertheless, females of many animals appear to increase the possibility of mating with compatible males; mate with more than one male; they are polyandrous it also permits pollen competition and therefore selection (e.g. Bateman, 1948; Birkhead and Mller, 1998 for a for pollen quality. Polyandry also increases the genetic review; Tregenza and Wedell, 2000; Portnoy et al., 2007; diversity of offspring (Ellstrand and Marshall, 1986; Frentiu and Chenoweth, 2008). As mating with multiple Armbruster and Rogers, 2004; Bernasconi et al., 2004; males incurs extra female costs (Cornell and Tregenza, Teixeira et al., 2009). As the number of ovules fertilized 2007), it is assumed that polyandry should either provide usually exceeds the number of mature seeds produced, direct or indirect advantages over mating with only one seeds with different paternity may go through a male. Indirect genetic advantages such as the avoidance postpollination selection process as well (Stephenson, of mating with incompatible males, opportunities for 1981; Wiens, 1984; Korbecka et al., 2002). Consequently, sperm competition, and cryptic female choice, may polyandry may have important implications for the explain the prevalence of polyandry in animals (see evolution of traits affecting the reception and donation of Cornell and Tregenza, 2007 for a review). pollen and the evolution of mating systems. Polyandry is prevalent in seed plants too in which Polyandry is assumed to have similar evolutionary female reproductive success appears to be resource implications in nonvascular embryophytes but informa- limited whereas male reproductive success is mainly tion concerning its occurrence and extent is scarce (Haig and Westoby, 1988; Haig and Wilczek, 2006). Moss Correspondence: Dr P Szo¨ve´nyi, Department of Biology, Duke University, gametophytes can be either bisexual (producing both 139 Biological Sciences Bldg., Box 90338, Durham, NC 27708, USA. E-mail: [email protected] archegonia with eggs and antheridia with sperm; Received 6 February 2009; revised 12 May 2009; accepted 4 June monoicous species) or unisexual (producing either 2009; published online 22 July 2009 archegonia or antheridia but not both; dioicous species). Multiple paternity in a dioicous moss P Szo¨ve´nyi et al 395 Unisexual gametophytes either outcross or undergo (multiple paternity)? We test this by genotyping the intergametophytic selfing (mating among haploid sibs maternal gametophyte plant and each attached sporo- from the same sporophyte), which is equivalent to phyte using microsatellite markers. As we then know the ‘selfing’ in seed plants. Bisexual gametophytes are also genotype of the female gamete (gametophyte) that bears capable of intragametophytic selfing (merging of ga- the diploid sporophyte, we can obtain the genotype of metes produced by the same genetic individual) result- the male gamete (gametophyte) by simple subtraction. ing in a fully homozygous sporophyte generation. (2) What is the average level of inbreeding in this Owing to this difference bisexual gametophytes are population and how do individual sporophytes vary in expected to rapidly purge recessive deleterious muta- the level to which they are inbred? (3) Is sporophyte tions through intragametophytic selfing, whereas more fitness related to the genotypes of the maternal and genetic load is assumed to accumulate in populations paternal gametophytes (that is, gametes); specifically, can of unisexual gametophytes (Eppley et al., 2007). This we detect evidence of inbreeding depression? We also difference in the amount of genetic load is proposed to assess the extent of spatial genetic structure in the have led to greater sporophytic inbreeding depression in population as this may promote fertilization among close species with unisexual compared with bisexual gameto- relatives. Finally, we discuss the possible roles and phytes (Taylor et al., 2007). If the severity of sporophytic evolutionary implications of polyandry in bryophytes. inbreeding depression is the primary force driving mating system evolution in bryophytes, species with unisexual gametophytes should be predominantly out- Materials and methods crossers. In contrast, species with bisexual gametophytes should predominantly inbreed because of the negligible Model species, plant material, and measuring of cost of inbreeding depression. However, natural popula- sporophyte traits tions of species with unisexual gametophytes are S. lescurii has unisexual (dioicous) gametophytes but is characterized by a mixed mating system (Eppley et al., characterized by frequent sporophyte production and 2007). In such a system polyandry with selective embryo occurrence of multiple sporophytes on individual female abortion may be able to decrease the realized costs of shoots. Young peatmoss sporophytes are surrounded by inbreeding and thus may explain the stability of mixed the perichaetial leaves of the female gametophyte and mating systems in species with unisexual gametophytes remain embedded for most of their life span. Unripe in face of inbreeding depression (Porcher and Lande, sporophytes are green and slowly turn black as they 2005). mature. In contrast to the majority of mosses, the One particularly important consequence of the freely peatmoss sporophyte consists of a spherical capsule swimming spermatozoids (as in mosses) is that fertiliza- and lacks a visible seta (stalk). When sporophytes are tion can only occur among individuals in close proximity mature the seta is replaced by an extension of the (McQueen, 1985; Wyatt, 1994; Crum, 2001; Bisang et al., maternal gametophyte, the pseudopodium, which en- 2004). This obviously limits the distance between ables successful spore dispersal by elevating the spor- successfully mating male and female gametophytes. ophyte above the female plant. As the pseudopodium is Furthermore, owing to the leptokurtic distribution of part of the female gametophyte, its elongation represents spore dispersal, a female plant is expected to be maternal investment. surrounded mainly by the haploid progeny of its S. lescurii samples were collected from a population in sporophyte (Longton, 1997; Sundberg, 2005; Gunnarsson Richmond County, North Carolina. The species forms a et al., 2007). Consequently, mating is likely to occur continuous mat at the margin of Lake Broadacres, with a among haploid siblings. Therefore, in the absence of a large proportion of gametophytes bearing multiple mechanism to distinguish among male gametes
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