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CONTENT - International Society of Zoological Sciences

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S11 ICZ2008 - Abstracts<br />

S12 - Transitions from clonal to sexual reproduction: key variations <strong>of</strong> a key process<br />

The function <strong>of</strong> sperm donating species to the evolution and<br />

persistence <strong>of</strong> unisexual salamanders (caudata: ambystomatidae)<br />

Jim Bogart<br />

Department <strong>of</strong> Integrative Biology, University <strong>of</strong> Guelph, N1G 2W1,<br />

Guelph, CANADA<br />

Unisexual salamanders in the genus Ambystoma are common<br />

around the Great Lakes region in eastern North America. Based on<br />

mitochondrial DNA sequences, they originated from a single<br />

hybridization event that involved a female that shared a common<br />

ancestor with Kentucky A. barbouri 2.4 to 3.9 million years ago.<br />

Individual unisexual salamanders reproduce by stealing sperm from<br />

donors <strong>of</strong> normally bisexual species so their reproductive mode is<br />

described as kleptogenesis. As many as 22 different diploid and<br />

polyploid unisexual biotypes are known and they all possess at least<br />

one A. laterale genome. One or more other nuclear genomes have<br />

been derived from sperm donors that may involve males <strong>of</strong> five<br />

distinct species. Sperm may serve only to stimulate egg<br />

development (gynogenesis) but can be incorporated to replace a<br />

nuclear genome or to elevate ploidy. Genome replacement is<br />

considered to be an essential evolutionary strategy to rectify meiotic<br />

problems that arise from recombinations and translocations among<br />

gynogenetic <strong>of</strong>fspring and to introduce novel genetic combinations<br />

that have a selective advantage. Molecular tools such as<br />

microsatellite DNA loci, expressed sequence tags (EST), and<br />

genomic in situ hybridization (GISH) can be used to identify the<br />

genomic constitution <strong>of</strong> individuals and to test hypotheses pertaining<br />

to the role <strong>of</strong> sperm donors in the co-evolution <strong>of</strong> the disparate<br />

cytoplasmic and nuclear components in unisexual salamanders.<br />

Bogart, J. 2003. Genetics and systematics <strong>of</strong> hybrid species. Pp.<br />

109-134, In D. M. Sever (edt.), Reproductive Biology and Phylogeny<br />

<strong>of</strong> Urodela. M/s Science Inc., Enfield NH.<br />

Bogart, J. P., K. Bi, J. Fu, D.W.A. Noble, and J. Niedzwieki. 2007.<br />

Unisexual salamanders (genus Ambystoma) present a new<br />

reproductive mode for eukaryotes. Genome 50: 119-136.<br />

Bi, K., J. P. Bogart, and J. Fu. 2008. Genealogical relationships <strong>of</strong><br />

unisexual salamanders <strong>of</strong> the genus Ambystoma inferred from<br />

intergenomic exchanges and 45S rDNA cytotypes. Chromosome<br />

Research 16: 275-289.<br />

Sex in asexuals: how occasional sex determines the fate <strong>of</strong><br />

parthenogenetic populations<br />

Thomas D'Souza<br />

Auf der Morgenstelle 26, 72076, Tuebingen, Germany<br />

Sex is expensive compared to asexual reproduction. Nevertheless<br />

sex is more widespread among multi-cellular organisms. Many<br />

theories have been developed to solve this so-called paradox <strong>of</strong> sex<br />

and to clarify why sex is advantageous. Particularly when combining<br />

these theories, the benefits <strong>of</strong> sex seem to outweigh the costs.<br />

Consequently, asexuals are usually considered as evolutionary<br />

dead-ends. This ra! ises the question whether asexual systems are<br />

always truly clonal or whether they have cryptic forms <strong>of</strong> sexuality<br />

that enhances their viability. This is important as, at least<br />

theoretically, a limited amount <strong>of</strong> sexuality is sufficient to compensate<br />

for the long-term costs <strong>of</strong> clonality. Here, the advantages <strong>of</strong><br />

occasional sex will be demonstrated using the planarian flatworm<br />

Schmidtea polychroa as an example. Parthenogenetic S. polychroa<br />

are simultaneous hermaphrodites, which require sperm from a<br />

partner in order to trigger embryo development (pseudogamy). As<br />

parthenogens are hermaphroditic, they produce fertile sperm and do<br />

not rely on sexual sperm donors. Parthenogenesis in S. polychroa is<br />

not always strictly clonal, but sometimes contains occasional, sexual<br />

process. This mixed reproduction mode may combine the benefits <strong>of</strong><br />

sex and asex and increases the survival <strong>of</strong> parthenogenetic<br />

populations in two different ways. Firstly, occasional sex promotes<br />

the investment in male reproductive organs <strong>of</strong> parthenogens, which<br />

is essential for rare sex among parthenogens.<br />

- 44 -<br />

Secondly, occasional sex in parthenogenetic S. polychroa leads to<br />

an increase in fitness variance at the local population level, which in<br />

turn increases mean fitness, which can be interpreted as a sign!<br />

ature <strong>of</strong> a more effective natural selection.<br />

Sexual reproduction in triploid forms <strong>of</strong> planarian Dugesia<br />

ryukyuensis<br />

Kazuya Kobayashi 1 , Hirotsugu Ishizu 1 , Motonori Hoshi 2 and Midori<br />

Matsumoto 1<br />

1<br />

Biosciences and Informatics, Keio University, Hiyoshi 3-14-1,<br />

Kouhoku-ku, Yokohama, 223-8522, Japan.<br />

2<br />

University <strong>of</strong> the Air, Wakaba 2-11, Mihama-ku, Chiba, 261-8586,<br />

Japan.<br />

Triploidy has generally been considered to be an evolutionary dead<br />

end due to problems <strong>of</strong> chromosomal pairing and segregation during<br />

meiosis. Thus, the formation <strong>of</strong> tetraploids and diploids from triploid<br />

types is a rare phenomenon. In the planarian Dugesia ryukyuensis,<br />

both fissiparous and oviparous triploids occur in nature. It is believed<br />

that oviparous triploids reproduce via pseudogamy (spermdependent<br />

parthenogenesis). We have successfully induced<br />

experimental sexualization <strong>of</strong> fissiparous triploids in D. ryukyuensis.<br />

Following sexualization, the triploids develop hermaphroditic gonads<br />

and other reproductive organs and begin reproducing by copulation<br />

instead <strong>of</strong> fission. In the present study, we demonstrated that<br />

inbreeding <strong>of</strong> the triploid planarian D. ryukyuensis resulted in both<br />

diploid and triploid <strong>of</strong>fspring in nature. In the triploids <strong>of</strong> D.<br />

ryukyuensis, chiasmata between homologous chromosomes were<br />

observed in both female and male germ lines. This result suggests<br />

that both diploid and triploid <strong>of</strong>fspring <strong>of</strong> this species are produced<br />

bisexually by zygotic fusion between sperm and eggs. Hence, this<br />

phenomenon may be a novel mechanism in planarian for escaping<br />

the triploid state.<br />

Males in a thelytokous strain <strong>of</strong> tardigrade<br />

Atsushi C. Suzuki<br />

Department <strong>of</strong> Biology, Keio University, Hiyoshi, Yokohama 223-<br />

8521, Japan<br />

Tardigrades are generally gonochoristic and propagate by sexual<br />

reproduction. A number <strong>of</strong> tardigrade species exhibit secondary<br />

sexual characters, with males in some Eutardigrada being<br />

distinguished by modified claws. Many moss-dwelling tardigrades,<br />

however, have thelytokous mode <strong>of</strong> parthenogenesis. One <strong>of</strong> such<br />

tardigrade is Milnesium tardigradum. Males in this species with<br />

modified claws are usually rare and many populations appear to<br />

have only parthenogenetic reproduction. The presence <strong>of</strong> males in<br />

small numbers, being greatly different from the sex ratio <strong>of</strong> 1:1, had<br />

not been considered as heterogony so far. Instead, it was interpreted<br />

to be the result <strong>of</strong> mixed populations that have parthenogenetic and<br />

amphimictic reproduction. Here the author reports that males have<br />

emerged at a very low frequency in a thelytokous strain <strong>of</strong> Milnesium<br />

cf. tardigradum, which has been maintained since 2000. Some<br />

individuals <strong>of</strong> this strain had modified claws characteristic <strong>of</strong> males<br />

on the first pair <strong>of</strong> legs. A small testis filling with many spermatozoa<br />

was observed in one <strong>of</strong> such specimen. Although copulation has not<br />

been observed yet, an interesting activity <strong>of</strong> a male against a female<br />

appeared to be a sexual behaviour. The frequency <strong>of</strong> the emergence<br />

<strong>of</strong> males in the cultured strain is so low that any environmental factor<br />

that may generate males has not been determined yet. It is unknown<br />

if these males could actually function as males in reproduction.<br />

However, they might show some possibility <strong>of</strong> genetic exchange<br />

among the clonal populations.

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