Effects of Pollen and Resources on Seed Number and Other Fitness Components in Amelanchier Arborea (Rosaceae: Maloideae) 1
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Amer. J. Bot. 75(9): 1275-1285. 1988. EFFECTS OF POLLEN AND RESOURCES ON SEED NUMBER AND OTHER FITNESS COMPONENTS IN AMELANCHIER ARBOREA (ROSACEAE: MALOIDEAE) 1 DAVID L. GORCHOV2 Department of Biology, University of Michigan, Ann Arbor, Michigan 48109 ABSTRACT Most Amelanchier arborea flowers have 10 ovules, but the number of filled (embryo-con taining) seeds per fruit is usually less than 10 and is highly variable within each individual plant. Because fruit developmental time correlates with seed number, this variation in seed number results in asynchronous fruit ripening. Field experiments tested whether seed number was pollen- or resource-limited. Manipulation of resources in the shoot at the time of fruit initiation by defoliation, girdling, fruit thinning, or foliar feeding had no significant effect on seed number per fruit, although fruit set and seed weight were affected. Supplemental cross pollination also had no demonstrable effect on seed number. Most ovules that do not become filled seeds are visible as small "undeveloped seeds"; these are not necessarily aborted seeds as this was also the fate of ovules in unpollinated carpels. Alternative hypotheses for the deter mination of seed number are proposed and discussed. THE NUMBER OF SEEDS that mature per fruit is few developing seeds are more likely to abort variable in many plant species and has several than those with many seeds (see reviews by important ecological, evolutionary, and hor Stephenson, 1981; Stephenson and Bertin, ticultural implications. Seed number has been 1983). By aborting fruits with low seed num studied as a component ofyield (or fitness) for bers, plants may improve the average quality its importance to both the adjustment of re of their offspring, either by eliminating fruits productive output under varying environmen resulting from self-fertilization (Darwin, 1876, tal conditions (Wilbur, 1977; Primack, 1978; pp. 398-400) or in which there has been little Wyatt, 1981; Schemske and Pautler, 1984; Ste pollen competition for ovules (Lee, 1984). phenson, 1984; Garwood and Horvitz, 1985; Seed number can affect seed dispersal in sev Marshall, Levin, and Fowler, 1985, 1986; Ga eral ways. Seed number is correlated with ma len and Weger, 1986) and the improvement of ture fruit size in many wild (Willson and agricultural crop yield (Olsson, 1960; Adams, Schemske, 1980; Bookman, 1983; Garwood 1967; Harper, 1977). In addition to its direct and Horvitz, 1985; Gorchov, 1985; Bronstein, contribution to total plant reproductive out 1986) as well as cultivated (Crane, 1964, p. put, seed number per fruit often influences the 303; Wareing and Phillips, 1978, p. 123) proportion of flowers that develop into fruits species; dispersers sometimes discriminate (fruit set). For several plant species, fruits with among fruits on the basis ofsize (Wheelwright, 1985). I Received for publication 22 June 1987; revision ac The ratio ofpulp weight to seed weight usu cepted 8 December 1987. I am grateful to K. Childs, J. Larsen, L. Whittingham, ally decreases with increasing seed number(e.g., B. Crespi, and K. Olsson for helping with field work, W. Willson and Schemske, 1980; Janzen, 1982). Frasch and J. Palmer for use of equipment, and D. Bay Because pulp is reward and seed is ballast to for photography. C. Campbell examined ovules for signs fruit-eating animals that disperse seeds, pulp: of apomictic seed development. M. Cruzan, L. Nooden, seed ratio should influence dispersers' prefer J. Hancock, and E. Hanson advised me on experimental methods. J. Bronstein, G. Estabrook, B. Rathcke, D. Gold ences (Howe and Vande Kerckhove, 1980; berg, B. Barnes, W. A. Robinson, and an anonymous re Herrera, 1981a, b). Similarly, seed number in viewer provided useful comments on earlier drafts ofthis fluences dispersal distance of some wind-dis manuscript. R. Kushler, G. Estabrook, and R. Bush aided persed species via its effects on mass/volume in data analysis. The Museum of Zoology, University of Michigan, provided access to the E. S. George Reserve ratio (Casperand Wiens, 1981; Augspurgerand and two ESGR scholarships. Additional support was pro Hogan, 1983; Augspurger, 1986). vided by Rackham Block grants from the University of Seed number may also affect the number of Michigan. This work represents part ofa dissertation sub seedlings germinating in one place, and there mitted in partial fulfillment ofthe requirements for a Ph.D. at the University of Michigan. fore sibling competition (Casper and Wiens, , Current address: Dept. of Biology, Princeton Univer 1981). Predispersal seed predators may prefer sity, Princeton, NJ 08544. fruits with more seeds (Herrera, 1984); where 1275 1276 AMERICAN JOURNAL OF BOTANY [Vol. 75 they don't have such a preference, spreading Cantrall (1943), and Cooper (1958). All Ame seeds few to a fruit reduces seed loss (Delph, lanchier arborea trees chosen for this study 1986). were from a dense population in oak-hickory Seed numbercan also affect dispersal through forest within 15 m ofa 270 m stretch ofa dirt its effects on ripening phenology. Within in road that runs along the top ofan esker (ridge). dividual plants ofsome species, the number of seeds is strongly negatively correlated with the duration of fruit development and with rip Study species-i-Amelanchier arborea (Michx. ening date (Aalders and Hall, 1961; Dempsey f.) Fern. (Rosaceae: Maloideae), Downy Ser and Boynton, 1965; Brewerand Dobson, 1969; viceberry, is a large shrub to small tree com Varga and Bruinsma, 1976; Lee and Bazzaz, mon in dry and mesic sites in eastern North 1982; Gorchov, 1985). For these plants, vari America (Fernald, 1950; Barnes and Wagner, ance in seed number results in ripening that is 1981). Species identifications were made using much more asynchronous than flowering (Gor Voss (1985); voucher specimens were depos chov, 1985). Thompson and Willson (1979) ited at the University ofMichigan Herbarium. proposed that the asynchronous ripening of Amelanchierarborea is one ofthe first shrubs temperate, summer-fruiting, bird-dispersed to bloom in the spring (late April--early May) plants is an adaptation to avoid satiation of at the study site; the white flowers open syn seed dispersal agents. chronously (90% offlowers on each plant open In the course oftesting Thompson and Will in two to five days, Gorchov, 1987) and are son's (1979) hypothesis, I became interested visited primarily by small bees (mostly An in the proximate (mechanistic) causes ofasyn drenidae and Halictidae) but also by other Hy chronous ripening. In the case ofAmelanchier menoptera, Diptera, and Coleoptera. arborea, a summer-ripening small tree with Flower morphology is similar to that de bird-dispersed seeds, flower-fruit interval is scribed for A. laevis (Campbell et al., 1985) and highly negatively correlated with the number A. alnifolia (Olson and Steeves, 1982): the gy offilled seeds (Gorchov, 1985). Hence, a prox noecium consists of five carpels each contain imate explanation for asynchronous ripening ing two ovules (hence a potential of 10 seeds in this species requires an understanding of per fruit), as in other Maloideae. The five styles why seed number varies among fruits. The im appear fused but are distinct ("pseudo-syn portance of resources, pollination, and seed carpous" cf. Carr and Carr, 1961) and diverge predation in the determination ofseed number at the stigmas, enabling separate pollination of raises the larger question of which of these individual carpels (Fig. 1). The inflorescence factors usually limits total plant reproductive is an elongate panicle and the terminal flower output (Bierzychudek, 1981; Stephenson, 1981; usually opens first. Inflorescences contain four Rathcke, 1983; Bawa and Webb, 1984). Snow to eleven flowers, with variability in flower (1986) showed that seed number is related to number relatively low within individual plants pollination intensity primarily in species with (SO = 0.7-2.2). very many seeds per fruit. Other studies have The fruits (pomes) are juicy, purple, and demonstrated a relationship between seed about 1 em in diameter when fully ripe. The number and resources (reviewed in Marshall number offully developed seeds per fruit ranges et al., 1986). from oneto 10, and the balance ofthe 10 ovules One objective ofthis study was to determine develop into small or empty "seeds" that have whether manipulation of resources in a shoot seed coats but no embryo. Amelanchier ar at the time offruit initiation affected fruit set, borea is the first species to ripen fleshy fruits seed number, or seed weight in A. arborea. I each summer in southeastern Michigan; fruits also tested whether addition of xenogamous ripen throughout June and early July. Fruit pollen to unbagged flowers increased fruit set ripening is relatively asynchronous within trees; or seed number compared to open-pollinated 90% of fruits ripen within 15-27 days (Gor controls. In addition I attempted to determine chov, 1987). Fruits are eaten by a variety of whether "undeveloped seeds" were aborted or birds and mammals; at the study site the prin unfertilized ovules. cipal seed disperser is the Cedar Waxwing, Bombycilla cedrorum. Chipmunks and other MATERIALSANDMETHODS-Studysite-Field rodents are seed predators (Robinson, 1986), work was done at the E. S. George Wildlife as are larvae oftortricid and geometrid moths. Reserve, Livingston County, Michigan, a re search area maintained by the University of Shoot-resource manipulations-Ten plants Michigan. Climate