Pollen size-number trade-off and pollen-pistil relationships in Pedicularis (Orobanchaceae)

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1 Plant Syst. Evol. 247: (2004) DOI /s Pollen size-number trade-off and pollen-pistil relationships in Pedicularis (Orobanchaceae) C.-F. Yang and Y.-H. Guo College of Life Sciences, Wuhan University, China Received April 15, 2003; accepted February 18, 2004 Published online: June 11, 2004 Ó Springer-Verlag 2004 Abstract. Current patterns of floral design in Pedicularis must have undergone an evolutionary process of interacting among components of floral traits, and then formed internal relationships among these traits. To detect such correlations, which may provide insight to understand flower evolution, 40 Pedicularis species representing all corolla types of the genus were studied. Results show that, interspecifically, pollen size correlates negatively with pollen number, but positively with pistil length. This suggests that plants evolve an optimal pollen size, which balances the advantages of large pollen size for gametophytic competition against the fecundity disadvantages of fewer pollen grains. In contrast to sex allocation theory, this study does not find a trade-off, but an interspecific positive correlation between pollen and ovule number. This is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the lack of negative correlation between male and female function. Key words: Evolution, ovule number, Pedicularis, pistil length, pollen number, pollen size, sex allocation. A central assumption underlying many arguments of plant reproductive evolution is that a genetically based negative correlation exists between module number and resource investment per module (Burd 1999). It is then expected that a pollen size-number trade-off is frequent in nature. Actually, both intra- and interspecific pollen size-number trade-off were found in several studies (intraspecific: Stanton and Young 1994, Vonhof and Harder 1995, Worley and Barrett 2000, Sarkissian and Harder 2001, but see Stanton and Preston 1986, Aguilar et al. 2002; interspecific: Mione and Anderson 1992, Vonhof and Harder 1995, but see Cruden and Miller-Ward 1981, Plitmann and Levin 1983, Knudsen and Olesen 1993, Aguilar et al. 2002). Such a negative relationship has been viewed as a consequence of the subdivision of limited resources in the plant (Vonhof and Harder 1995). As one component of integrated floral design, pollen size cannot evolve independently, but it coevolves with other floral traits, e.g. style length (Sarkissian and Harder 2001). A pistil length increase will necessarily result in an increase of pollen size; otherwise pollen tubes would simply fail to reach the ovules. Thus the resulting sexual selection will tend to create a positive correlation between pollen size and style length (Plitmann and Levin 1983). This principle has been supported by many studies (Plitmann and Levin 1983, Harder 1998, Roulston et al. 2000, Torres

2 178 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships 2000, Sarkissian and Harder 2001, Aguilar et al. 2002, but see Cruden and Miller-Ward 1981, Cruden and Lyon 1985). Besides the pollen size-number trade-off, there also should be a trade-off in resource allocation between male function (pollen) and female function (ovule) based on sex allocation theory (Charlesworth and Charlesworth 1981, Charnov 1982). Darwin (1877) had presented the law of compensation that reduced investment into one reproductive function may be compensated by the greater resource availability for the other reproductive function. Consequently, a negative correlation between investment in male and female function is expected in cosexual species (Stearns 1992). However, most recent studies have detected positive correlations, and only few have reported negative correlations (reviewed by Koelewijin and Hunscheid 2000, Campbell 2000). Campbell (2000) explained the positive correlation and considered both positive and negative relationship between resources allocation into the two sexual functions to be reasonable. Pedicularis, a large hermaphrodite genus whose species display substantial variation in floral design from a narrow genetic basis (Macior 1995), is a monophyletic group (Yang et al. 2003). More than 350 Pedicularis species have been recorded from China, with most being restricted to the Southwest of the country (Hong 1983, Yang et al. 1998). The Chinese Himalaya is regarded as the centre of origin and evolution of Pedicularis, where species with all corolla types of the genus are present (Li 1951, Yang et al. 2003). Pedicularis species provide ideal material to study the following objectives: (1) Working on a small monophyletic group Lycieae, which is pollinated by similar pollinators, Aguilar et al. (2002) did not find an interspecific pollen size-number trade-off. The lack of a trade-off was explained as a result of the similar constraints experienced by the species and/or because constraints are operating at the plant level and not the at the species level. Although Pedicularis species are pollinated exclusively by bumblebees (Macior and Tang 1997, Macior et al. 2001), they display relatively high interspecific variation in pollen size (Harder 1998, Yang et al. 2002). Does pollen size correlate negatively with pollen number per flower at the species level in Pedicularis species with similar pollinators? (2) Pedicularis species display a very high variation in style length (pistil length) (Yang et al. 1998). What the evolutionary significance of the long style is remains unclear up to now (Macior and Tang 1997, Yang et al. 2002). If pistil length and pollen size covary (Sarkissian and Harder 2001), is there a functional relation between the two traits? (3) If pollen number correlates with pollen size and displays a high level of variation in Pedicularis species, according to the theory of sex allocation, does there exist a trade-off relation between pollen and ovule number? Materials and methods A total of 40 Pedicularis species (Table 1) representing all corolla types in the genus were collected from the native habitat from 2000 to All species were found in the Northwest of Yunnan province and West of Sichuan province in Southwest China. Voucher specimens are preserved in the Wuhan University Herbarium (WH), P. R. China. Studied flowers were collected randomly from plants in natural populations for each species. Sample sizes of each variable are shown in Table 1. Mature fully-opened flowers for further study in the laboratory were picked randomly during the peak blooming period in each species and quickly fixed in FAA solution constituted of formalin (37 40%), acetic acid and alcohol (50%) at a ratio of 5: 6: 89 by volume. Pistil length was considered an approximation to the maximum distance that a pollen tube must grow to fertilize an ovule (Torres 2000). The distance corresponded to the length between the stigmatic tip and the base of the ovary, since Pedicularis has basal placentation. Mature flowers with undehisced anthers were used to measure pollen number. Anthers were dissected and placed into vials containing 0.3 ml 0.7 M mannitol solution colored with aceto carmine to stain the cytoplasm. Pollen was counted in a haemocytometer after shaking thoroughly. The

3 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships 179 Table 1. Floral traits of 40 Pedicularis species (mean±sd). For each species, no less than 20 individual plants were used for measurement of pistil length, 10 for pollen grain number, 20 for ovule number, and 30 for pollen grain volume Species Pistil length (cm) N>35 Pollen grain number/flower N>10 Ovule number/ ovary N>25 Pollen grain volume (lm 3 ) N>80 P. anas Maxim ± ± ± ± P. axillaris Franch ± ± ± ± P. batangensis Bureau 3.93 ± ± ± ± et Franch. P. confertiflora Prain 2.70 ± ± ± ± P. cyathophylla Franch ± ± ± ± P. cymbalaria Bonati 3.01 ± ± ± ± P. davidii Franch ± ± ± ± P. debilis Franch ± ± ± ± P. densispica Franch ± ± ± ± P. dichotoma Bonati 2.62 ± ± ± ± P. dolichocymba Hand.-Mazz ± ± ± ± P. dolichoglossa H. L. Li 2.53 ± ± ± ± P. dunniana Bonati 2.71 ± ± ± ± P. elwesii Hook. f ± ± ± ± P. geosiphon H. Sm ± ± ± ± et P. C. Tsoong P. gracilicaulis H. L. Li 2.82 ± ± ± ± P. gruina Franch ± ± ± ± P. integrifolia Hook. f ± ± ± ± P. kansuensis Maxim ± ± ± ± P. lachnoglossa Hook. f ± ± ± ± P. latituba Bonati 6.93 ± ± ± ± P. likiangensis Franch ± ± ± ± P. longiflora Rudolph var. longiflora 7.84 ± ± ± ± P. longiflora Rudolph var. tubiformis (Koltz.) 7.44 ± ± ± ± P. C. Tsoong P. macrosiphon Franch ± ± ± ± P. megalochila H. L. Li 3.66 ± ± ± ± P. monbeigiana Bonati 2.39 ± ± ± ± P. mussoti Franch ± ± ± ± P. polyodonta H. L. Li 6.94 ± ± ± ± P. princeps Bureau et Franch ± ± ± ± P. rex C. B. Clarke ssp ± ± ± ± lipskyana (Bonati) P. C. Tsoong P. rex C. B. Clarke ssp. rex 3.87 ± ± ± ± ovary was carefully dissected out of each flower and placed in a drop of water on a microscope slide. The entire placenta with attached ovules was removed via a longitudinal slit in the ovary wall. The ovules were carefully removed from the placenta and spread in the drop of water to be counted at 40 magnification under a dissecting microscope. Size of pollen grains was estimated by

4 180 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships Table 1 Species (continued) Pistil length (cm) N>35 Pollen grain number/flower N>10 Ovule number /ovary N>25 Pollen grain volume (lm 3 ) N>80 P. rhinanthoides Schrenk ex 4.78 ± ± ± ± Fisch. et Mey. ssp. Labellata (Jacq.) P. C. Tsoong P. rhinanthoides Schrenk ex 3.98 ± ± ± ± Fisch. et Mey. ssp. rhinanthoides P. rhodotricha Maxim ± ± ± ± P. roylei Maxim ± ± ± ± P. siphonantha Don 6.94 ± ± ± ± P. stenocorys Franch ± ± ± ± P. szetschuanica Maxim ± ± ± ± P. tenera H. L. Li 1.79 ± ± ± ± P. trichoglossa Hook. f ± ± ± ± P. tricolor Hand.-Mazz. ssp ± ± ± ± tricolor P. tricolor Hand.-Mazz. ssp ± ± ± ± lophocentra H. L. Li P. urceolata P. C. Tsoong 3.99 ± ± ± ± measuring the diameter of the polar and equatorial axes of the ellipsoidal grains from dehisced anthers. Measurements were made with an ocular micrometer at 400. The volume of single pollen grains was estimated by the formula ppe 2 /6 (Aguilar et al. 2002), where P is the polar axis diameter and E is the equatorial axis diameter. We used the SAS program (SAS Institute 1998) for all statistical analyses. Correlation analysis and tests of significance were used to determine relationships between all variables for all the study species. Coefficients of variation (CV) were calculated for pollen grain volume, pollen grain number per flower. Means of original data±sd are given in the text and tables. Results and discussion Mean values of pollen grain volume, pollen grain number per flower, pistil length, and ovule number per flower for all the study species are summarized in Table 1. The variables vary in a sixfold to twenty-five-fold range in the studied species. Statistical analysis based on the studied 40 species shows that pollen grain number per flower correlates negatively with pollen grain volume (r ¼ )0.424, P ¼ 0.004). An interspecific pollen size-number trade-off exists in Pedicularis species (Fig. 1a). In addition, a very strong interspecific linear positive correlation was found between mean pistil length and mean pollen grain volume (r ¼ 0.705, P < , Fig. 1b). Considering resource allocation to male versus female function, a positive relationship exists between pollen grain number and ovule number (r ¼ 0.446, P ¼ 0.002, Fig. 1c). Coefficient of variation (CV) analysis shows that at both the level of genus and species, pollen grain number per flower varies considerably more than pollen grain volume and ovule number per flower (Table 2). Pollen size-number trade-off. Interspecific inverse relationships between size and number of pollen grains have also been documented in some other studies (Mione and Anderson 1992, Vonhof and Harder 1995). Subdivision of limited resources is ascribed to resulting in the pollen size-number trade-off (Vonhof and Harder 1995). Because resources available for the male function are limited, any increase in

5 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships r=-0.424** a r=0.705**** b Pollen size (µm 3 ) Pollen size (µm 3 ) Pollen grain number Pistil length (cm) 100 r=0.446** c Ovule number Pollen grain number Fig. 1. Relationship of floral traits in Pedicularis. Significance level: **** < 0:0001; *** 0:0001 < P < 0:0001; ** 0:001 < P < 0:01; 0:01 < P < 0:05

6 182 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships Table 2. Coefficient of variation (CV) of pollen grain number, and pollen grain volume in Pedicularis, data from 40 species Variable CV for whole genus a Mean CV of all species b Pollen grain number per flower Pollen grain volume **** a Data of the two variables from all 40 species are listed and calculated to get the CV value b The average of CV values for all 40 species, significance level: ****P< pollen size may lead to a trade-off with pollen number (Vonhof and Harder 1995, Sarkissian and Harder 2001). Pollen size-number trade-off also suggests that the evolution of pollen number in response to a specific pollination environment must be influenced by the physiological and ecological advantages of different pollen sizes (see also Vonhof and Harder 1995, Harder 1998). On studying interspecific size-number trade-off in pollen, the phylogenetic relatedness and pollinator type must be taken into account for the examined species. Several interspecific studies did not find a negative correlation between pollen size and number (Cruden and Miller-Ward 1981, Plitmann and Levin 1983, Aguilar et al. 2002). However, reanalysis of data of Cruden and Miller-Ward (1981) by removing the phylogenetic relatedness (family), Vonhof and Harder (1995) found a clear inverse relation between pollen size and pollen number per flower. The absence of size-number trade-off in the study by Plitmann and Levin (1983) on 170 species in Polemoniaceae may be because the diversity of pollinator types was not taken into account (Vonhof and Harder 1995). Pedicularis, a monophyletic genus with a uniform pollinator type (see Introduction), displays a pollen size-number trade-off based on analysis data of 40 species (Fig. 1a). In contrast, Aguilar et al. (2002) did not find a trade-off between pollen size and number in a small monophyletic group Lycieae whose species have all the same pollinators. However, if the closest relatives are compared in this data set, an interspecific pollen size-number trade-off can also be found. We reanalyzed their data of five varieties of Lycium chilense (Table 1 in Aguilar et al. 2002) and found an inverse relation between pollen volume and pollen number (r=)0.899, P<0.05). This suggests that interspecific pollen size-number trade-off may evolve from the interaction between resource allocation to male and female function among closely related groups with a uniform pollination system. Pollen size is less variable than most other floral traits (Cresswell 1998). Comparison of CV values between pollen size and pollen grain number per flower in this study supports the argument very well. CV value of pollen size is smaller than that of pollen number at both level of genus and species (Table 2). A similar pattern was also observed by Vonhof and Harder (1995), and Aguilar et al. (2002). This result suggests that there is a stronger resilience to variations of reproductive resources of pollen size compared with pollen number (Vonhof and Harder 1995, and references therein). Pollen size, rather than pollen number may be the primary target of natural selection (see also Sarkissian and Harder 2001). Pistil length and pollen size. Considering the functional relationship between the energy storage capacity of pollen grains and the stigma-ovule distance, Baker and Baker (1982) suggested that species with long pistils tend to have large pollen grains to provide enough energy for pollen tube germination and growth. Besides this study, a positive correlation between pistil length and pollen grain size has also been detected in a large number of unrelated plant taxa (Baker and Baker 1982, Plitmann and Levin 1983, Williams and Rouse

7 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships , Harder 1998, Bigazzi and Selvi 2000, Roulston et al. 2000, Torres 2000, Sarkissian and Harder 2001, Aguilar et al. 2002, but see Cruden and Miller-Ward 1981, Cruden and Lyon 1985). Positive correlation between pistil length and pollen size may result from a widespread gametic-phase disequilibrium that arises from nonrandom fertilization success of large pollen in pistils with long style (Sarkissian and Harder 2001). Mulcahy (1983) showed that increased style length intensifies male-to-male competition and selects for faster pollen tube growth rate. Moreover, there is evidence that larger pollen grains can give rise to faster growing pollen tubes (Lord and Eckard 1984). The positive correlation between pollen size and pistil length in Pedicularis (Fig. 1b) indicates that large pollen grains in species with long pistil afford the essential prerequisite for pollen competition by possessing enough substance to provide energy for pollen tube growth to pass the long style and fertilize the ovules. The presence of correlations between pollen grain volume and pistil length, and pollen grain number per flower in Pedicularis suggests that pollen size could not have evolved independently (see also Sarkissian and Harder 2001). Torres (2000) suggests that three aspects of the reproductive processes, viz., resource allocation to male function, pollination, and post-pollination processes must be related to the evolution of pollen size. He also mentions that post-pollination events may play a more important role in pollen size evolution compared to resource allocation to male function and pollination events. Provided that pollen grain volume has probably undergone stronger natural selection than pollen grain number and the pollinators of the species in Pedicularis are similar, pollen size evolution in the genus must be correlated with post-pollination processes to a greater extent. The finding of a significant correlation between pistil length and pollen grain volume supports this argument, and also suggests that for Pedicularis species, pollen size may have co-evolved with pistil length (see also Aguilar et al. 2002). Pollen and ovule number. The basic premise of sex allocation theory is that there should be a trade-off in resource allocation between male and female function, and that plants should distribute resources to these functions to obtain optimal fitness (Charlesworth and Charlesworth 1981, Charnov 1982, Morgan 1992). However, the majority of recent studies about this topic have revealed a wide intraspecific within-flower positive genetic correlation between male and female function (Campbell 1992, 1997; Mazer 1992, O Neill and Schmitt 1993, Agren and Schemske 1995, Ashman 1999, Burd 1999, Koelewijn and Hunscheid 2000). The first and one of the very few studies demonstrating a negative correlation between pollen and ovule number was an artificial selection study in Spergularia marina by Mazer et al. (1999). Similar to Ashman (1999), Burd (1999) and Koelewijn and Hunscheid (2000), we also found a positive correlation between pollen and ovule number in 40 species of Pedicularis (Fig. 1c). In addition, our study provides a new case by finding an interspecific trade-off. Such interspecific studies are somewhat neglected and may give new insight to understand the different opinions about resource allocation to male versus female function (see also Koelewijn and Hunscheid 2000). The lack of a negative, but presence of a positive correlation between resource investment to male and female function is not surprising (Campbell 2000), it may in part be ascribed to genetic variation for resource acquisition (Campbell 2000, Koelewijn and Hunscheid 2000), and perhaps to the fact that flowers can draw on different resource pools for male and female parts (Ashman 1994, Campbell 2000). In summary, our finding that pollen size-number trade-off and positive relationship between pollen size and pistil length suggest that pollen size cannot evolve independently. Plants evolve to create an optimal pollen size that balances the advantages of large pollen size for gametophytic competition against the fecundity disadvantages of fewer pollen grains.

8 184 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships The result that pollen number correlates positively with ovule number is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the lack of negative correlation between male and female function. This work was supported by the State Key Basic Research and Development Plan, China (Grant No. G ) to YHG. The authors would like to thank Peter K. Endress and two anonymous reviewers for providing critical comments and helpful suggestions, Qing-Feng Wang, Jing-Yuan Wang and Jin-Ming Chen for their helpful suggestions. Shi-Guo Sun, Jing Xia, and Qian Yu are thanked for their assistance in both the field work and laboratory phases of the project. References Agren J., Schemske D. W. (1995) Sex allocation in the monoecious herb Begonia semiovata. Evolution 49: Aguilar R., Bernardello G., Galetto L. (2002) Pollenpistil relationships and pollen size-number tradeoff in species of the tribe Lycieae (Solanaceae). J. Plant Res. 115: Ashman T.-L. (1994) A dynamic perspective on the physiological cost of reproduction in plants. Amer. Nat. 144: Ashman T.-L. (1999) Determinants of sex allocation in a gynodioecious wild strawberry: implications for the evolution of dioecy and sexual dimorphism. J. Evol. Biol. 12: Baker H. G., Baker I. (1982) Starchy and starchless pollen in the Onagraceae. Ann. Missouri Bot. Gard. 69: Bigazzi M., Selvi F. (2000) Stigma form and surface in the tribe Boragineae (Boraginaceae); micromorphological diversity, relationships with pollen, and systematic relevance. Canad. J. Bot. 78: Burd M. (1999) Flower number and floral components in ten angiosperm species: an examination of assumptions about trade-offs in reproductive evolution. Biol. J. Linn. Soc. 68: Campbell D. R. (1992) Variation in sex allocation and floral morphology in Ipomopsis aggregata (Polemoniaceae). Amer. J. Bot. 79: Campbell D. R. (1997) Genetic correlation between biomass allocation to male and female functions in a natural population of Ipomopsis aggregata (Polemoniaceae). Heredity 79: Campbell D. R. (2000) Experimental tests of sexallocation theory in plants. Trends Ecol. Evol. 15: Charlesworth B., Charlesworth D. (1981) Allocation to the male and female function in hermaphrodites. Biol. J. Linn. Soc. 15: Charnov E. L. (1982) The theory of sex allocation. Princeton Univ. Press, Princeton, NJ. Cresswell J. E. (1998) Stabilizing selection and the structural variability of flowers within species. Ann. Bot. 81: Cruden R. W., Lyon D. L. (1985) Correlations among stigma depth, style length, and pollen grain size: do they reflect function or phylogeny? Bot. Gaz. 146: Cruden R. W., Miller-Ward S. (1981) Pollen-ovule ratio, pollen size, and the ratio of stigmatic area to the pollen-bearing area of the pollinator: an hypothesis. Evolution 35: Darwin C. (1877) Different forms of flowers on plants of the same species. John Murray, London. Harder L. D. (1998) Pollen-size comparisons among animal-pollinated angiosperms with different pollination characteristics. Biol. J. Linn. Soc. 64: Hong D.-Y. (1983) The distribution of Scrophulariaceae in the Holarctic in reference to the floristic relationships between Eastern Asia and Eastern North America. Ann. Missouri Bot. Gard. 70: Knudsen J. T., Olesen J. M. (1993) Buzz-pollination and patterns in sexual traits in North European Pyrolaceae. Amer. J. Bot. 80: Koelewijn H. P., Hunscheid M. P. H. (2000) Intraspecific variation in sex allocation in hermaphroditic Plantago coronopus (L.) J. Evol. Biol. 13: Li H.-L. (1951) Evolution in the flowers of Pedicularis. Evolution 5: Lord E. M., Eckard (1984) Incompatibility between the dimorphic flowers of Collomia grandiflora, a cleistogamous species. Science 223: Macior L. W. (1995) Pedicularis, a valuable information resource for plant conservation. In: Sihag R. C. (ed.) Pollination biology. Rajendra Scientific Pulishers, Hisar, pp

9 C.-F. Yang and Y.-H. Guo: Pollen size-number trade-off and pollen-pistil relationships 185 Macior L. W., Tang Y. (1997) A preliminary study of the pollination ecology of Pedicularis in the Chinese Himalaya. Plant Species Biol. 12: 1 7. Macior L. W., Tang Y., Zhang J.-C. (2001) Reproductive biology of Pedicularis (Scrophulariaceae) in the Sichuan Himalaya. Plant Species Biol. 16: Mazer S. J. (1992) Environmental and genetic sources of variation in floral traits and phenotypic gender in wild radish: consequences for natural selection. In: Wyatt R. (ed.) Ecology and evolution of plant reproduction. Chapman & Hall, New York, pp Mazer S. J., Delesalle V. A., Neal P. R. (1999) Responses of floral traits to selection on primary sexual investment in Spergularia marina: the battle between the sexes. Evolution 53: Mione T., Anderson G. J. (1992) Pollen-ovule ratios and breeding system evolution in Solanum section Basarthrum (Solanaceae). Amer. J. Bot. 79: Morgan M. (1992) The evolution of traits influencing male and female fertility in outcrossing plants. Amer. Nat. 139: Mulcahy D. L. (1983) Models of pollen tube competition in Geranium maculatum. In: Real L. (ed.) Pollination biology. Academic Press, London, pp O Neill P., Schmitt J. (1993) Genetic constraints on the independent evolution of male and female reproductive characters in the tristylous plant Lythrum salicaria. Evolution 47: Plitmann U., Levin D. A. (1983) Pollen-pistil relationships in the Polemoniaceae. Evolution 37: Roulston T., Cane J. H., Buchmann S. L. (2000) What governs protein content of pollen: pollinator preferences, pollen-pistil interactions, or phylogeny? Ecol. Monogr. 70: Sarkissian T. S., Harder L. D. (2001) Direct and indirect responses to selection on pollen size in Brassica rapa L. J. Evol. Biol. 14: SAS. (1998) SAS/STAT user s guide. Cary. NC: SAS Institute. Stanton M. L., Preston R. E. (1986) Pollen allocation in wild radish: variation in pollen grain size and number. In: Mulcahy D. L., Mulcahy B., Ottaviano E. (eds.) Biotechnology and ecology of pollen. Springer, New York, pp Stanton M. L., Young H. J. (1994) Selecting for floral character associations in wild radish, Raphanus sativus L. J. Evol. Biol. 7: Stearns S. C. (1992) The evolution of life histories. Oxford University Press, Oxford. Torres C. (2000) Pollen size evolution: correlation between pollen volume and pistil length in Asteraceae. Sex. Plant Reprod. 12: Vonhof M. J., Harder L. D. (1995) Size-number trade-offs and pollen production by papilionaceous legumes. Amer. J. Bot. 82: Williams E. G., Rouse J. L. (1990) Relationships of pollen size, pistil length and pollen tube growth rates in Rhododendron and their influence on hybridization. Sex. Plant Reprod. 3: Worley A. C., Barrett S. C. H. (2000) Evolution of floral display in Eichhornia paniculata (Pontederiaceae): direct and correlated responses to selection on flower size and number. Evolution 54: Yang C.-F., Guo Y.-H., Gituru R. W., Sun S.-G. (2002) Variation in stigma morphology How does it contribute to pollination adaptation in Pedicularis (Orobanchaceae)? Plant Syst. Evol. 236: Yang F.-S., Wang X.-Q., Hong D.-Y. (2003) Unexpected high divergence in nrdna ITS and extensive parallelism in floral morphology of Pedicularis (Orobanchaceae). Plant Syst. Evol. 240: Yang H., Holmgren N. H., Mill R. R. (1998) Pedicularis L. In: Raven P. H. (ed.) Flora of China (18). Science Press, Beijing, pp Address of the authors: Chun-Feng Yang and You-Hao Guo ( College of Life Sciences, Wuhan University, Wuhan , China.

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