Sexual selection: an evolutionary force in plants?
IO SKOGSMYR,
ÅSA LANKINEN,
IO SKOGSMYR
Department of Theoretical Ecology, Ecology Building, Lund University, 223 62 Lund, Sweden (E-mail: [email protected] )
Search for more papers by this authorÅSA LANKINEN
Department of Theoretical Ecology, Ecology Building, Lund University, 223 62 Lund, Sweden (E-mail: [email protected] )
Search for more papers by this author
IO SKOGSMYR,
ÅSA LANKINEN,
IO SKOGSMYR
Department of Theoretical Ecology, Ecology Building, Lund University, 223 62 Lund, Sweden (E-mail: [email protected] )
Search for more papers by this authorÅSA LANKINEN
Department of Theoretical Ecology, Ecology Building, Lund University, 223 62 Lund, Sweden (E-mail: [email protected] )
Search for more papers by this authorABSTRACT
Sexual selection has traditionally been used to explain exaggerated sexual traits in male animals. Today the concept has been developed and various other sexually related traits have been suggested to evolve in the same manner. In nearly all new areas where the theory of sexual selection has been applied, there has been an intense debate as to whether the application is justified. Is it the case that some scientists are all too ready to employ fashionable ideas? Or are there too many dogmatic researchers refusing to accept that science develops and old ideas are transformed? Maybe the controversies are simply a reflection of the difficulty of defining a theory under constant re-evaluation. Thus, we begin by summarizing the theory of sexual selection in order to assess the influence of sexual selection on the evolution of plant morphology. We discuss empirical findings concerning potentially affected traits. Although we have tried to address criticisms fairly, we still conclude that sexual selection can be a useful tool when studying the evolution of reproductive traits in plants. Furthermore, by including the evidence from an additional kingdom, a fuller understanding of the processes involved in sexual selection can be gained.
VII. REFERENCES
- Adams, S., Vinkenoog, R., Spielman, M., Dickinson, H. G. & Scott, R. J. (2000). Parent-of-origin effects on seed development in Arabidopsis thaliana require DNA methylation. Development 127, 2493–2502.
- Aizen, M. A., Searcy, K. B. & Mulcahy, D. L. (1990). Among- and within-flower comparisons of pollen tube growth following self- and cross-pollinations in Dianthus chinensis (Caryophyllaceae). American Journal of Botany 77, 671–676.
- Alatalo, R. V., Mappes, J. & Elgar, M. A. (1997). Heritabilities and paradigm shifts. Nature 385, 402–403.
- Andersson, M. (1986). Evolution of condition-dependent sex ornaments and mating p: sexual selection based on viability differences. Evolution; International Journal of Organic Evolution 40, 804–826.
- Andersson, M. (1994). Sexual Selection. Princeton University Press, Princeton .
- Andersson, M. & Iwasa, Y. (1996). Sexual selection. Trends in Ecology and Evolution (Amsterdam) 11, 53–58.
- Arathi, H. S., Ganeshaiah, K. N., Uma Shaanker, R. & Hegde, S. G. (1996). Factors affecting embryo abortion in Syzygium cuminii (L.) skeels (Myrtaceae). International Journal of Plant Sciences 157, 49–52.
- Armbruster, W. S. (1996). Evolution of floral morphology and function: an integrative approach to adaptation, constraint, and compromise in Dalechampia (Euphorbiaceae). In Floral Biology (eds. D. G. Lloyd and S. C. H. Barrett), pp. 241–272. Chapman & Hall, New York .
10.1007/978-1-4613-1165-2_9 Google Scholar
- Armbruster, W. S., Martin, P., Kidd, J., Stafford, R. & Rogers, D. G. (1995). Reproductive significance of indirect pollen-tube growth in Dalechampia (Euphorbiaceae). American Journal of Botany 82, 51–56.
- Arnold, J. S. (1994). Is there a unifying concept of sexual selection that applies to both plants and animals American Naturalist 144 (suppl.), 1–12.
- Bakker, T. C. M. (1993). Positive genetic correlation between female preference and preferred male ornament in sticklebacks. Nature 363, 255–257.
- Balmford, A. (1991). Mate choice on leks. Trends in Ecology and Evolution (Amsterdam) 6, 87–91.
- Barnes, H., Barnes, M. & Klepal, W. (1977). Studies on the reproduction of Cirripedes. I. Introduction: copulation, release of oocytes, and formation of the egg lamellae. Journal of Experimental Marine Biology and Ecology 27, 195–218.
- Bateman, A. J. (1948). Intra-sexual selection in Drosophila. Heredity 2, 349–368.
- Bensch, S. & Hasselquist, D. (1991). Evidence for an active female choice in a polygynous warbler. Animal Behaviour 44, 301–312.
- Beletsky, L. D. (1983). Agressive and pair-bond maintenance songs of female red-winged blackbirds (Agelaius phoeniceus). Z. Tierpsycol 62, 47–54.
- Bertin, R. I. (1988). Paternity in plants. In Plant Reproductive Ecology: Patterns and Strategies (eds. J. Lovett Doust and L. Lovett Doust), pp. 30–59. Oxford University Press, New York .
- Bertin, R. I. (1990). Effects of pollination intensity in Campsis radicans. American Journal of Botany 77, 178–187.
- Björkman, T. (1995). The effect of pollen load and pollen grain competition on fertilization success and progeny performance in Fagopyrum esculentum. Euphytica 83, 47–52.
- Björkman, T., Samimy, C. & Pearson, K. J. (1995). Variation in pollen performance among plants of Fagopyrum esculenyum. Euphytica 82, 235–240.
- Bond, W. J. & Maze, K. E. (1999). Survival costs and reproductive benefits of floral display in a sexually dimorphic dioecious shrub, Leucadendron xanthoconus. Evolutionary Ecology 13, 1–18.
- Bookman, S. S. (1984). Evidence for selective fruit production in Asplepias. Evolution; International Journal of Organic Evolution 38, 72–86.
- Brewbaker, J. L. & Majunder, S. K. (1961). Cultural studies of the pollen population effect and the self-incompatibility inhibition. American Journal of Botany 48, 457–464.
- Broyles, S. B. & Wyatt, R. (1990). Paternity analysis in a natural population of Asclepias exaltata: multiple paternity, functional gender, and the ‘pollen-donation hypothesis’. Evolution; International Journal of Organic Evolution 44, 1454–4689.
- Broyles, S. B. & Wyatt, R. (1995). A reexamination of the pollen-donation hypothesis in an experimental population of Asclepias exalta. Evolution 49, 89–99.
- Campbell, D. R. (1989). Measurements of selection in a hermaphroditic plant: Variation in male and female pollination success. Evolution; International Journal of Organic Evolution 43, 318–334.
- Casper, B. B. (1988). Evidence for selective embryo abortion in Cryptantha flava. American Naturalist 132, 318–326.
- Charlesworth, B. (1987). The heritability of fitness. In Sexual Selection: Testing the alternatives (eds. J. W. Bradbury and M. B. Andersson), pp. 21–40. Wiley, New York .
- Charlesworth, D., Schemske, D. W. & Sork, V. L. (1987). The evolution of plant reproductive characters: sexual versus natural selection. In Sexual Selection: Testing the Alternatives (eds. J. W. Bradbury and M. B. Andersson), pp. 317–335. Wiley, New York .
- Charnov, E. L. (1979). Simultaneous hermaphroditism and sexual selection. Proceedings of the National Academy of Sciences of the United States of America 76, 2480–2484.
- Cheung, A. Y. (1995). Pollen-pistil interactions in compatible pollination. Proceedings of the National Academy of Sciences of the United States of America 92, 3077–3080.
- Cheung, A. Y., Wang, H. & WU, H.-M. (1995). A floral transmitting tissue-specific glycoproten attracts pollen tubes and stimulates their growth. Cell 82, 383–393.
- Choi, J.-S. & Friedman, W. E. (1991). Development of the pollen tube of Zamia furfuracea (Zaniaceae) and its evolutionary implications. American Journal of Botany 78, 544–560.
- Cruzan, M. & Barrett, S. C. (1993). Contribution of cryptic incompatibility to the mating system of Eichhornia paniculata (Pontederiaceae). Evolution; International Journal of Organic Evolution 47, 925–934.
- Cruzan, M. B. (1990a). Pollen-pollen and pollen-style interaction during pollen tube growth in Erythronium grandiflorum (Liliaceae). American Journal of Botany 77, 116–122.
- Cruzan, M. B. (1990b). Variation in pollen size, fertilization ability and postfertilization siring ability in Erythronium grandiflorum. Evolution; International Journal of Organic Evolution 44, 843–856.
- Cruzan, M. B. (1993). Analysis of pollen-style interactions in Petunia hybrida; the determination of variance in male reproductive success. Sexual Plant Reproduction 6, 275–281.
- Cruzan, M. B., Neal, P. R. & Willson, M. F. (1988). Floral display in Phyla incisa: conseqenses for male and female reproductive success. Evolution; International Journal of Organic Evolution 3, 505–515.
- Cruzan, M. B. & Thomson, J. D. (1997). Effects of pre-disperal selection on offspring growth and survival in Erythronium grandiflorum. Journal of Evolutionary Biology 10, 295–314.
- Cunningham, E. J. A. & Birkhead, T. R. (1998). Sex roles and sexual selection. Animal Behaviour 56, 1311–1321.
- Dahl, Å. E. & Fredrikson, M. (1996). The timetable for development of maternal tissues sets the stage for male genomic selection in Betula pendula (Betulaceae). American Journal of Botany 83, 895–902.
- Dajoz, I., Till-Bottraud, I. & Gouyon, P.-H. (1991). Evolution of pollen morphology. Science 253, 66–68.
- Darwin, C. (1871). The Descent of Man and Selection in Relation to Sex. John Murray, London .
- De Jong, T. J. & Klinkhamer, P. G. L. (1994). Plant size and reproductive success through female and male function. Ecology 82, 399–402.
- Delph, L. F., Johannsson, M. H. & Stephenson, A. G. (1997). How environmental factors affect pollen performance: ecological and evolutionary perspectives. Ecology 78, 1632–1639.
- Delph, L. F., Weinig, C. & Sullivan, K. (1998). Why fastgrowing pollen tubes give rise to vigorous progeny: the test of a new mechanism. Proceedings of the Royal Society of London Series B: Biological Sciences 265, 935–939.
- Devlin, B., Clegg, J. & Ellstrand, N. C. (1992). The effect of flower production on male reproductive success in wild radish populations. Evolution; International Journal of Organic Evolution 46, 1030–1042.
- Dominey, W. J. (1983). Sexual selection, Additive Genetic Variance and the ‘Phenotypic Handicap’. Journal of Theoretical Biology 101, 495–502.
- Douglas, K. L. & Cruden, R. W. (1994). The reproductive biology of Anemone candensis (Ranunculaceae): breeding system and facilitation of sexual selection. American Journal of Botany 81, 314–321.
- Eberhard, W. G. (1996). Female Control: Sexual Selection by Cryptic Female Choice. Princeton University Press, Princeton .
10.1515/9780691207209 Google Scholar
- Eckhart, V. M. (1993). Do hermaphrodites of gymnodioecious Phacelia linearis (Hydrophyllaceae) trade off seed production to attract pollinators Biological Journal of the Linnean Society 50, 47–63.
- Emms, S. K., Stratton, D. A. & Snow, A. A. (1997). The effect of influorescence size on male fitness: experimental tests in the andromonoecious lily, Zigadenus paniculatus. Evolution; International Journal of Organic Evolution 51, 1481–1489.
- Falconer, D. S. & Mackay, T. F. C. (1996). Introduction to Quantitative Genetics. 4th ed. Longman, London .
- Fenster, C. B. (1991). Effect of male pollen donor and female seed parent on allocation of resources to developing seeds and fruits in Chamaecrista fasciculata (Leguminosae). American Journal of Botany 78, 13–23.
- Fenster, C. B. & Sork, V. L. (1988). Effect of crossing distance and male parent on in vivo pollen tube growth in Chamaecrista fasciculata. American Journal of Botany 75, 1898–1903.
- Fishbein, M. & Venable, L. D. (1996). Evolution of inflorescence: theory and data. Evolution; International Journal of Organic Evolution 50, 2165–2177.
- Fisher, R. A. (1958). The Genetical Theory of Natural Selection. Clarendon Press, Oxford .
- Friedman, W. (1995). Organismal duplication, inclusive fitness theory, and altruism: understanding the evolution of endosperm and the angiosperm reproductive syndrome. Proceedings of the National Academy of Sciences of the United States of America 92, 3913–3917.
- Furlow, B. (1999). Flower power. New Scientist 161, 22–26.
- Galen, C. (1992). Pollen dispersal dynamics in an alpine wildflower, Polemonium viscusum. Evolution; International Journal of Organic Evolution 46, 1043–1051.
- Gambier, R. M. & Mulcahy, D. L. (1996). The association between pollen size and Renner complex in Oenothera villaricae and O. picensis ssp. Picensis and their hybrids: evidence for preanthesis pollen competition. Theoretical and Applied Genetics 92, 140–144.
- Ganeshaiah, K. N. & Uma Shaanker, R. (1988). Regulation of seed number and female incitation of mate competition by a pH-dependent proteinaceous inhibitor of pollen grain germination in Leucaena leucocephala. Oecologia 75, 110–113.
- Grafen, A. (1990). Sexual selection unhandicapped by the Fisher process. Journal of Theoretical Biology 144, 473–516.
- Grant, V. (1995). Sexual selection in plants: pros and cons. Proceedings of the National Academy of Sciences of the United States of America 92, 1247–1250.
- Hamilton, W. D. (1980). Sex versus non-sex versus parasite. Oikos 35, 282–290.
- Hamilton, W. D. & Zuk, M. (1982). Heritable true fitness and bright birds: a role for parasites Science 218, 384–387.
- Harder, L. D. & Barrett, S. C. H. (1995). Mating cost of large floral displays in hermaphrodite plants. Nature 373, 512–515.
- Harder, L. D. & Thomson, J. D. (1989). Evolutionary options for maximizing pollen dispersal of animal-pollinated plants. American Naturalist 133, 323–344.
- Härdling, R. & Nilsson, P. (2001). A model of triploid endosperm evolution driven by parent-offspring conflict. Oikos 92, 417–423.
- Havens, K. (1994). Clonal repeatability of in vitro pollen tube growth rates in Oenothera organensis (Onagraceae). American Journal of Botany 81, 161–165.
- Havens, K. & Delph, L. F. (1996). Differential seed maturation uncouples fertilization and siring success in Oenothera organensis (Onagraceae). Heredity 76, 623–632.
- Herrero, M. (1983). Factors affecting fruit set in ‘Agua de Aranjuez’ pear. Acta Hortic 139, 91–96.
10.17660/ActaHortic.1983.139.12 Google Scholar
- Herrero, M. & Hormaza, J. I. (1996). Pistil strategies controlling pollen tube growth. Sexual Plant Reproduction 9, 343–347.
- Hill, J. P. & Lord, E. M. (1986). Dynamics of pollen tube growth in the wild radish, Raphanus raphanistrum (Brassicaceae). I. Order of fertilization. Evolution; International Journal of Organic Evolution 40, 1328–1333.
- Hormaza, J. I. & Herrero, M. (1994). Gametophytic competition and selection. In Genetic Control of Self-incompatibility and Reproductive Development in Flowering Plants (eds. E. G. Williams, A. E. Clarke and R. B. Knox), pp. 372–400. Kluwer, Dordrecht .
10.1007/978-94-017-1669-7_18 Google Scholar
- Hormaza, J. I. & Herrero, M. (1996a). Dynamics of pollen tube growth under different competition regimes. Sexual Plant Reproduction 9, 153–160.
- Hormaza, J. I. & Herrero, M. (1996b). Male gametophytic selection as a plant breeding tool. Scientia Horticulturae 65, 321–333.
- Houle, D. (1992). Comparing evolvability and variability of quantitative traits. Genetics 130, 195–204.
- Hrdy, S. B. (1981). The Woman that Never Evolved. Harvard University Press.
- Iwasa, Y., Pomiankowski, A. & Nee, S. (1991). The evolution of costly mate p. II. The ‘handicap’ principle. Evolution; International Journal of Organic Evolution 45, 1431–1442.
- Johannsson, M. H. & Stephenson, A. G. (1997). Effects of pollination intensity on the vigor of the sporophytic and gametophytic generation of Cucurbita texana. Sexual Plant Reproduction 10, 236–240.
- Johannsson, M. H. & Stephenson, A. G. (1998). Effects of temperature during microsprogenesis on pollen performance in Cucurbita pepo L. (Cucurbitaceae). International Journal of Plant Sciences 159, 616–626.
- Johnston, M. O. (1993). Tests of two hypotheses concerning pollen competition in a self-compatible, long-styled species (Lobelia cardinalis: Lobeliaceae). American Journal of Botany 80, 1400–1406.
- Johnstone, R. A. (1995). Sexual selection, honest advertisement and the handicap principle: reviewing the evidence. Biological Reviews of the Cambridge Philosophical Society 70, 1–65.
- Kirkpatrick, M. & Ryan, M. J. (1991). The evolution of mating p and the paradox of the lek. Nature 350, 33–38.
- Kobayashi, S., Inoue, K. & Kato, M. (1997). Evidence of pollen transfer efficiency as the natural selection factor favoring a large corolla of Campanula punctata pollinated by Bombus diversus. Oecologia 111, 535–542.
- Kodric-Brown, A. & Brown, J. H. (1984). Truth in advertising: the kinds of traits favored by sexual selection. American Naturalist 124, 309–323.
- Lande, R. (1976). The maintenance of genetic variability by mutation in a polygenic character with linked loci. Genetical Research 26, 221–235.
- Lande, R. (1981). Models of speciation by sexual selection on polygenic traits. Proceedings of the National Academy of Sciences of the United States of America 78, 3721–3725.
- Landi, P. & Frascaroli, E. (1988). Pollen-style interactions in Zea mays L. In Sexual Reproduction in Higher Plants (eds. M. Cresti, P. Gori and E. Pacini), pp. 315–320. Springer, Berlin , Heidelberg , New York .
10.1007/978-3-642-73271-3_50 Google Scholar
- Lank, D. B., Smith, C. M., Hanotte, O., Burke, T. & Cooke, F. (1995). Genetic polymorphism for alternative mating bahaviour in lekking male ruff Philomachus pugnax. Animal Behaviour 378, 59–62.
- Lankinen, Å. (2001). In vitro pollen competitive ability in Viola tricolor: temperature and pollen donor effects. Oecologia 128, 492–498.
- Lankinen, Å. (2000). Effects of soil phosphorus on in vitro pollen competitive ability and sporophytic traits clones of Viola tricolor. International Journal of Plant Sciences 161, 885–893.
- Lankinen, Å. & Skogsmyr, I. (2002). Pollen competitive ability: the effect of proportion in two-donor crosses. Evolutionary Ecology Research in press.
- Lankinen, Å. & Skogsmyr, I. (2001a). Evolution of pistil length as a choice mechanism for pollen quality. Oikos 92, 81–90.
- Lankinen, AI. & Skogsmyr, I. (2001b). The effect of pollen competition on maintenance of variation in fertilization ability. Oikos 93, 459–469.
- Lau, T.-C. & Stephenson, A. G. (1993). Effects of soil nitrogen on pollen production, pollen grain size, and pollen performance in Cucurbita pepo (Cucurbitaceae). American Journal of Botany 80, 763–768.
- Lau, T.-C. & Stephenson, A. G. (1994). Effects of soil phoshorus on pollen production, pollen size, pollen phosphorus content, and the ability to sire seeds in Cucurbita pepo (Cucurbitaceae). Sexual Plant Reproduction 7, 215–220.
- Leach, C. R., Renfrey, A. C. P. & Mayo, O. (1993). Quantitatively determined self-incompatibility, IV. Pollination and seed set in Borago officinalis. Theoretical and Applied Genetics 85, 755–760.
- Lee, T. D. (1984). Patterns of fruit maturation: a gametophyte competition hypothesis. American Naturalist 123, 427–432.
- Lovett Doust, J. (1990). Botany agonistes: on phytocentrism and plant sociobiology. Trends in Ecology and Evolution (Amsterdam) 4, 121–133.
- Lovett Doust, J. & Lovett Doust, L. (1988). Sociobiology of plants: an emerging synthesis. In Plant Reproductive Ecology, Patterns and Strategies (eds. J. Lovett Doust and L. Lovett Doust), pp. 5–29. Oxford University Press, Oxford .
- Lyons, E. E., Waser, N. M., Price, M. V., Antonovics, J. & Motten, A. F. (1989). Sources of variation in plant reproductive success and implications for concepts of sexual selection. American Naturalist 134, 409–433.
- Majerus, M. E. N., O'Donald, P., Kearns, P. W. E. & Ireland, H. (1986). Genetics and evolution of female choice. Nature 321, 164–167.
- Marshall, D. L. (1988). Postpollination effects on seed paternity: mechanisms in addition to microgametophyte competition operate in wild radish. Evolution; International Journal of Organic Evolution 42, 1256–1266.
- Marshall, D. L. (1998). Pollen donor performance can be consistent across maternal plants in wild radish (Raphanus sativus, Brassicaceae): a necessary condition for the action of sexual selection. American Journal of Botany 85, 1389–1397.
- Marshall, D. L., Avritt, J. J., Shaner, M. & Saunders, R. L. (2000). Effects of pollen load size and composition on pollen donor performance in wild radish, Raphanus sativus (Brassicaceae). American Journal of Botany 87, 1619–1627.
- Marshall, D. L. & Diggle, P. K. (2001). Mechanisms of differential pollen donor performance in wild radish, Raphanus sativus (Brassicaceae). American Journal of Botany 88, 242–257.
- Marshall, D. L. & Ellstrand, N. C. (1986). Sexual selection in Raphanus sativus: experimental data on nonrandom fertilization, maternal choice, and consequences of multiple paternity. American Naturalist 127, 446–461.
- Marshall, D. L. & Folsom, M. W. (1991). Mate choice in plants: an anatomical to population perspective. Annual Review of Ecology and Systematics 22, 37–63.
- Marshall, D. L., Folsom, M. W., Hatfield, C. & Bennett, T. (1996). Does interference competition among pollen grains occur in wild radish Evolution; International Journal of Organic Evolution 50, 1842–1848.
- Marshall, D. L. & Fuller, O. S. (1994). Does nonrandom mating among wild radish plants occur in the field as well as in the greenhouse American Journal of Botany 81, 439–445.
- Marshall, D. L. & Oliveras, D. M. (1990). Is regulation of mating within branches possible in wild radish (Raphanus sativus L.) Functional Ecology 4, 619–627.
- Marshall, D. L. & Whittaker, K. L. (1989). Effects of pollen donor quality in wild radish, Raphanus sativus. American Journal of Botany 76, 1081–1088.
- Maynard Smith, J. (1978). The handicap principle-a comment. Journal of Theoretical Biology 70, 251–252.
- Mazer, S. J. (1987a). Maternal investment and male reproductive success in angiosperms: parent-offspring conflict or sexual selection Biological Journal of the Linnean Society 30, 115–133.
- Mazer, S. J. (1987b). Parental effects on seed development and seed yield in Raphanus raphanistrum: implications for natural and sexual selection. Evolution; International Journal of Organic Evolution 41, 355–371.
- McKenna, M. A. & Mulcahy, D. L. (1983). Ecological aspects of gametophytic competition in Diantus chinensis. In Pollen: Biology and Implications for Plant Breeding (eds. D. L. Mulcahy and E. Ottaviano), pp. 419–424. Elsevier, New York .
- Meager, T. R. (1991). Analysis of paternity within a natural population of Chamaelirium luteum. II Patterns of male reproductive success. American Naturalist 137, 738–752.
- Melser, C., Rademaker, M. C. J. & Klinkhammer, P. G. L. (1997). Selection on pollen donors by Echium vulgare (Boraginaceae). Sexual Plant Reproduction 10, 305–312.
- Merilä, J. & Sheldon, B. C. (2000). Lifetime reproductive success and heritability in nature. American Naturalist 155, 301–310.
- Mitchell, R. J. & Marshall, D. L. (1998). Nonrandom mating and sexual selection in adester mustard: an experimental approach. American Journal of Botany 85, 48–55.
- Mitchell, R. J., Shaw, R. G. & Waser, N. M. (1988). Pollinator selection, quantitative genetics, and predicted evolutionary responses of floral traits in Penstemon centranthifolius (Scrophulariaceae). International Journal of Plant Sciences 159, 331–337.
- Mitchell, R. L. (1997). Effects of pollen quantity on progeny vigor: evidence from the desert mustard Lesquella fendleri. Evolution; International Journal of Organic Evolution 51, 1679–1684.
- Moore, T. & Haig, D. (1991). Genomic imprinting in mammalian development: a parental tug-of-war. Trends in Genetics 7, 45–49.
- Morgan, M. T. (1992). The evolution of traits influencing male and female fertility in outcrossing plants. American Naturalist 139, 1022–1051.
- Morgan, M. T. (1994). Models of sexual selection in hermaphrodites, especially plants. American Naturalist 144 (suppl.), 100–125.
- Morgan, M. T. & Schoen, D. J. (1997). Selection on reproductive characters: floral morphology in Asclepias syriaca. Heredity 79, 433–441.
- Mulcahy, D. L. (1971). A correlation between gametophytic characters in Zea mays L. Science 171, 1155–1156.
- Mulcahy, D. L. (1979). The rise of the angiosperms: a genecological factor. Science 206, 20–23.
- Mulcahy, D. L., Curtis, P. S. & Snow, A. A. (1983). Pollen competition in a natural population. In Handbook of experimental pollination biology (eds. C. E. Jones and R. J. Little), pp. 330–337. Van Nostrand Reinhold, New York .
- Mulcahy, D. L. & Mulcahy, G. B. (1985). Ovarian influence on pollen tube growth, as indicated by the semivivo technique. American Journal of Botany 72, 1078–1080.
- Mulcahy, D. L. & Mulcahy, G. B. (1987). The effects of pollen competition. American Scientist 75, 44–50.
- Mulcahy, D. L., Mulcahy, G. B. & Searcy, K. B. (1992). Evolutionary genetics of pollen competition. In Ecology and Evolution of Plant Reproduction (ed. R. Wyatt), pp. 25–36. Chapman and Hall, New York .
- Mulcahy, D. L., Sari-Gorla, M. & Mulcahy, G. B. (1996). Pollen selection - past, present and future. Sexual Plant Reproduction 9, 353–356.
- Mulcahy, G. B., Mulcahy, D. L. & Pfahler, P. L. (1982). The effect of delayed pollination in Petunia hybrida. Acta Botanica Neerl 31, 97–103.
- Mutikainen, P. & Delph, L. F. (1995). Effects of herbivory on male reproductive success in plants. Oikos 75, 353–358.
- Namai, H. & Ohsawa, R. (1992). Evidence for non-random seed setting of ovules in the pod of the cruciferous plants Brassica juncea and Raphanus sativus. In Angiosperm Pollen and Ovules (eds. E. Ottaviano, D. L. Mulcahy, M. Sari Gorla and G. B. Mulcahy), pp. 429–434. Springer-Verlag, New York .
10.1007/978-1-4612-2958-2_70 Google Scholar
- Niesenbaum, R. A. (1996). Linking herbivory and pollination: defoliation and selective fruit abortion in Lindera benzoin. Ecology 77, 2324–2331.
- Niesenbaum, R. A. (1999). The effects of pollen load size and donor diversity on pollen performance, selective abortion, and progeny vigor in Mirabilis jalapa (Nyctaginaceae). American Journal of Botany 86, 261–268.
- Niesenbaum, R. A. & Casper, B. B. (1994). Pollen tube numbers and selective fruit maturation in Lindera benzoin. American Naturalist 144, 184–191.
- Niovi Jones, K. & Reithel, J. S. (2001). Pollinator-mediated selection on a flower color polymorphism in experimental populations of Antirrhinum (Scrophulariaceae). American Journal of Botany 88, 447–454.
- O'Donnell, M. E. & Bawa, K. S. (1993). Gamete selection and patterns of ovule and seed abortion. Current Science 65, 214–219.
- Ottaviano, E., Sari-Gorla, M. & Villa, M. (1988). Pollen competitive ability in maize: within population variability and response to selection. Theoretical and Applied Genetics 76, 601–608.
- Ottaviano, E. M., Sari-Gorla, M. & Arenari, I. (1983). Male gametophyte competitive ability in maize: selection and implications with regard to the breeding system. In Pollen: Biology and Implications for Plant Breeding (eds. D. L. Mulcahy and E. Ottaviano), pp. 367–374. Elsevier, NY .
- Palmer, T. M. & Zimmerman, M. (1994). Pollen competition and sporophyte fitness in Brassica campestris: does intense pollen competition result in individuals with better pollen Oikos 69, 80–86.
- Pasonen, H., Käpylä, M. & Pulkkinen, P. (2000). Effects of temperature and pollination site on pollen performance in Betula pendula Roth - evidence for genotype-environment interactions. Theoretical and Applied Genetics 100, 1108–1112.
- Pasonen, H. L., Pulkkinen, P. & Käpylaä, M. (2001). Do pollen donors with fastest-growing pollen tubes sire the best offspring in an anemophilous tree, Betula pendula (Betulaceae) American Journal of Botany 88, 854–860.
- Pasonen, H. L., Pulkkinen, P., Käpylä, M. & Blom, A. (1999). Pollen tube growth rate and seed siring success among Betula pendula clones. New Phytologist 143, 243–251.
- Partridge, L. (1983). Non-random mating in offspring fitness. In Mate Choice (ed. P. Bateson), pp. 227–256. Cambridge University Press, Cambridge .
- Pfahler, P. L. (1967). Fertilization ability of maize pollen grains. II. Pollen genotype, female sporophyte, and pollen storage interactions. Genetics 57, 513–521.
- Pfahler, P. L., Pereira, M. J. & Barnett, R. D. (1997). Genetic variation for in vitro sesame pollen germination and tube growth. Theoretical and Applied Genetics 95, 1218–1222.
- Philipp, M. & Hansen, T. (2000). The influence of plant and corolla size on pollen deposition and seed set in Geranium sanguineum (Geraniaceae). Nordic Journal of Botany 20, 129–140.
- Pleasants, J. M. (1981). Bumblebee response to variation in nectar availability. Ecology 62, 1648–1661.
- Pleasants, J. M. & Stephen, S. J. (1983). Nectar production rates of Asclepas quadrifolia: causes and consequences of individual variation. Oecologia 59, 232–238.
- Pomiankowski, A., Iwasa, Y. & Nee, S. (1991). The evolution of costly mate p. I. Fisher and biased mutation. Evolution; International Journal of Organic Evolution 45, 1422–1430.
- Pomiankowski, A. & Møller, A. P. (1995). A resolution of the lek paradox. Proceedings of the Royal Society of London Series B: Biological Sciences 260, 21–29.
- Potts, W. K. & Wakeland, E. K. (1993). Evolution of MHC genetic diversity: a tale of incest, persilence and sexual preference. Trends in Genetics 9, 408–412.
- Price, T. & Schluter, D. (1991). On the low heritability of lifehistory traits. Evolution; International Journal of Organic Evolution 45, 853–861.
- Queller, D. (1997). Pollen removal, paternity, and the male function of flowers. American Naturalist 149, 585–594.
- Queller, D. C. (1983). Sexual selection in a hermaphroditic plant. Nature 305, 706–707.
- Queller, D. C. (1987). Sexual selection in flowering plants. In Sexual Selection: Testing the Alternatives (eds. J. W. Bradbury and M. B. Andersson), pp. 165–179. Wiley, New York .
- Queller, D. C. (1994). Male-female conflict and parent- offspring conflict. American Naturalist 144 (suppl.), 84–99.
- Quesada, M., Bollman, K. & Stephenson, A. G. (1995). Leaf damage decreases pollen production and hinders pollen performance in Cucurbita texana. Ecology 76, 437–443.
- Quesada, M., Winsor, J. A. & Stephenson, A. G. (1993). Effects of pollen competition on progeny performance in a heterozygous cucurbit. American Naturalist 142, 694–706.
- Quesada, M. R., Winsor, J. A. & Stephenson, A. G. (1996). Effects of pollen selection on progeny vigor in a Cucurbita pepo × C. texana hybrid. Theoretical and Applied Genetics 92, 885–890.
- Rademaker, M. C. J. & De Jong, T. J. (1998). Effects of flower number on estimated pollen transfer in natural populations of three hermaphroditic species: an experiment with fluorescent dye. Journal of Evolutionary Biology 11, 623–641.
- Reynolds, J. D. & Gross, M. R. (1990). Costs and benefits of female mate choice: is there a lek paradox American Naturalist 136, 230–243.
- Rice, W. R. (1988). Heritable variation in fitness as a prerequisite for adaptive female choice: the effect of mutation- selection balance. Evolution; International Journal of Organic Evolution 42, 817–820.
- Richards, A. J. (1997). Plant Breeding Systems. Chapman and Hall, London .
10.1007/978-1-4899-3043-9 Google Scholar
- Richter, J. & Powels, S. B. (1993). Pollen expression of herbicide target site resistance genes in annual ryegrass (Lolium rigidum). Plant Physiology 102, 1037–1041.
- Ritchie, M. (1996). What is the ‘the paradox of the lek’ Trends in Ecology and Evolution (Amsterdam) 11, 175.
- Rocha, O. J. & Stephenson, A. G. (1991). Effects of nonrandom seed abortion on progeny performance in Phaseolus coccineus L. Evolution; International Journal of Organic Evolution 45, 1198–1208.
- Rowe, L. & Houle, D. (1996). The lek paradox and the capture of genetic variance by condition dependent traits. Proceedings of the Royal Society of London Series B: Biological Sciences 263, 1415–1421.
- Sanders, L. C. & Lord, E. M. (1989). Direct movement of latex particles in the gynoecia of three species of flowering plants. Science 243, 1606–1608.
- Sari-Gorla, M., Pé, M. E., Mulcahy, D. L. & Ottaviano, E. (1992). Genetic dissection of pollen competitive ability in maize. Heredity 69, 423–430.
- Sarr, A., Fraleigh, B. & Sandmeier, M. (1983). Aspects of reproductive biology in pearl millet, Pennisetum typhoides (Burm.) stapf and hubb. In Pollen: Biology and Implications for Plant Breeding (eds. D. L. Mulcahy and E. Ottaviano), pp. 381–388. Elsevier, New York .
- Schemske, D. W. & Fenster, C. (1983). Pollen-grain interactions in a neotropical Costus: effects of clump size and competitors. In Pollen: Biology and Implications for Plant Breeding (eds. D. L. Mulcahy and E. Ottaviano), pp. 405–410. Elsevier, New York .
- Schlichting, C. D., Stephenson, A. G., Small, L. E. & Winsor, J. A. (1990). Pollen loads and progeny vigour in Cucurbita pepo: the next generation. Evolution; International Journal of Organic Evolution 44, 1358–1372.
- Scott, R. J., Spielman, M., Bailey, J. & Dickinson, H. G. (1998). Parent-of-origin effects on seed development in Arabidopsis thaliana. Development 125, 3329–3341.
- Searcy, W. A. (1992). Song repertoire and mate choice in birds. American Zoologist 32, 71–80.
- Shaanker, R. U. & Ganeshaiah, K. N. (1997). Conflict between parent and offspring in plants: predictions, processes and evolutionary consequences. Current Science 72, 932–939.
- Skogsmyr, I. & Lankinen, A. I. (1999). Selection on pollen competitive ability in relation to stochastic factors influencing pollen deposition. Evolutionary Ecology Research 1, 971–985.
- Skogsmyr, I. & Lankinen, A. I. (2000). Female assessment of good genes in stylar tissue. Evolutionary Ecology Research 2, 965–979.
- Snow, A. A. (1986). Pollination dynamics in Epilobium canum (Onagraceae): consequences for gametophytic selection. American Journal of Botany 73, 139–151.
- Snow, A. A. (1990). Effects of pollen-load size and number of donors on sporophyte fitness in wild radish (Raphanus raphanistrum). American Naturalist 136, 742–758.
- Snow, A. A. (1991). Effects of pollen-load size on sporophyte competitive ability in two epilobium species. Am Midl Nat 125, 348–355.
- Snow, A. A. (1994). Postpollination selection and male fitness in plants. American Naturalist 144 (suppl.), 69–83.
- Snow, A. A. & Mazer, S. J. (1988). Gametophytic selection in Raphanus raphanistrum: a test for heritable variation in pollen competitive ability. Evolution; International Journal of Organic Evolution 42, 1065–1075.
- Snow, A. A. & Spira, T. P. (1991a). Differential pollen-tube growth rates and nonrandom fertilization in Hibiscus moscheutos (Malvaceae). American Journal of Botany 78, 1419–1426.
- Snow, A. A. & Spira, T. P. (1991b). Pollen vigour and the potential for sexual selection in plants. Nature 352, 796–797.
- Snow, A. A. & Spira, T. P. (1996). Pollen-tube competition and male fitness in Hibiscus moscheutos. Evolution; International Journal of Organic Evolution 50, 1866–1870.
- Spira, T. P., Snow, A. A. & Puterbaugh, M. N. (1996). The timing and effectiveness of sequential pollinations in Hibiscus moscheutos. Oecologia 105, 230–235.
- Spira, T. P., Snow, A. A., Whigham, D. F. & Leak, J. (1992). Flower visitation, pollen deposition, and pollen-tube competition in Hibiscus moscheutos (Malvaceae). American Journal of Botany 79, 428–433.
- Stanton, M. L. (1994). Male-male competition during pollination in plant populations. American Naturalist 144 (suppl.), 40–68.
- Stanton, M. L., Snow, A. A. & Handel, S. N. (1986). Floral evolution: attractiveness to pollinators increases male fitness. Science 232, 1625–1627.
- Stearns, S. C. (1992). The Evolution of Life Histories. Oxford University Press, New York .
- Stephenson, A. G. (1980). Fruit set, herbivory, fruit reduction, and the fruiting strategy of Catalpa speciosa (Bignoniaceae). Ecology 61, 57–64.
- Stephenson, A. G. (1992). The regulation of maternal investment in plants. In Fruit and seed production. Aspects of development, environmental physiology and ecology. Society for Experimental Biology Seminar Series 47 (eds. C. Marshall and J. Grace), pp. 151–172. Cambridge University Press, Cambridge .
- Stephenson, A. G. & Bertin, R. I. (1983). Male competition, female choice and sexual selection in plants. In Pollination Biology (ed. L. Real), pp. 109–149. Academic Press, London .
10.1016/B978-0-12-583980-8.50013-2 Google Scholar
- Stephenson, A. G. & Winsor, J. A. (1986). Lotus corniculatus regulates offspring quality through selective fruit abortion. Evolution; International Journal of Organic Evolution 40, 453–458.
- Sutherland, S. (1987). Why hermaphroditic plants produce many more flowers than fruits: Experimental tests with Agave mckelveyana. Evolution; International Journal of Organic Evolution 41, 750–759.
- Thomson, J. D. (1988). Effects of variation in influorescence size and floral rewards on the visitation rates of traplining pollinators of Aralia hispida. Evolutionary Ecology 2, 65–76.
- Thomson, J. D., Rigney, L. P., Karoly, K. M. & Thomson, B. A. (1994). Pollen viability, vigour, and competitive ability in Erythronium grandiflorum (Liliaceae). American Journal of Botany 81, 1257–1266.
- Travers, S. E. (1999). Environmental effects on components of pollen performance in Faramea occidentalis (L.) A. Rich. (Rubiaceae). Biotropica 31, 159–166.
- Travers, S. E. & Holtsford, T. P. (2000). Differential siring success of Pgi genotypes in Clarkia Ungulata (Onagraceae). American Journal of Botany 87, 1091–1098.
- Tregenza, T. & Wedell, N. (2000). Genetic compatibility, mate choice and patterns of parentage: Invited Review. Molecular Ecology 9, 1013–1027.
- Trivers, R. L. (1972). Parental investment and sexual selection. In Sexual Selection and the Descent of Man, 1871–1971 (ed. B. Campbell), pp. 136–179. Heinemann, London .
- Turner, G. F. (1995). The lek paradox resolved Trends in Ecology and Evolution (Amsterdam) 10, 473–474.
- Vaughton, G. & Ramsey, M. (1998). Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae). Oecologia 115, 93–101.
- Walsh, N. E. & Charlesworth, D. (1992). Evolutionary interpretations of differences in pollen tube growth rates. Quarterly Review of Biology 67, 19–37.
- Wedekind, C. (1994). Mate choice and maternal selection for specific parasite resistances before, during and after fertilization. Philosophical Transactions of the Royal Society of London Series B: Biological Sciences 346, 303–311.
- West-Eberhard, M. J. (1979). Sexual selection, social competition and speciation. Proceedings of the American Philosophical Society 123, 222–134.
- Wiens, D., Calvin, C. L., Wilson, C. A., Davern, C. I., Frank, D. & Seavey, S. R. (1987). Reproductive success, spontaneous embryo abortion, and genetic load in flowering plants. Oecologia 71, 501–509.
- Willson, M. F. (1979). Sexual selection in plants. American Naturalist 113, 777–790.
- Willson, M. F. (1990). Sexual selection in plants and animals. Trends in Ecology and Evolution (Amsterdam) 5, 210–214.
- Willson, M. F. (1994). Sexual selection in plants: perspective and overview. American Naturalist 144 (suppl.), 13–39.
- Willson, M. F. & Burley, N. (1983). Mate Choice in Plants: Tactics, Mechanisms and Consequences. Princeton University Press, Princeton .
- Wilson, P., Thomson, J. D., Stanton, M. L. & Rigney, L. P. (1994). Beyond floral batemania: gender biases in selection for pollination success. American Naturalist 143, 283–296.
- Winsor, J. A., Davis, L. E. & Stephenson, A. G. (1987). The relationship between pollen load and fruit maturation and the effect of pollen load on offspring vigor in Cucurbita pepo. American Naturalist 129, 643–656.
- Winsor, J. A., Peretz, S. & Stephenson, A. G. (2000). Pollen competition in a natural population of Cucurbita foetidissima (Cucurbitaceae). American Journal of Botany 87, 527–532.
- Young, H. J. & Stanton, M. L. (1990). Influence of environmental quality on pollen competitive ability in wild radish. Science 248, 1631–1633.
- Zahavi, A. (1975). Mate selection - a selection for a handicap. Journal of Theoretical Biology 53, 205–214.
- Zimmerman, M. (1988). Nectar production, flowering phenology, and strategies for pollination. In Plant Reproductive Ecology: Patterns and Strategies (eds. J. Lovett Doust and L. Lovett Doust), pp. 157–178. Oxford University Press, New York .
