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Life History Tradeâ•'Offs and Evidence for Hierarchical Resource

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Life History Tradeâ•'Offs and Evidence for Hierarchical Resource Plant Biology ISSN 1435-8603 RESEARCH PAPER Life history trade-offs and evidence for hierarchical resource allocation in two monocarpic perennials G.-X. Cao1,2 & A. C. Worley3 1 Department of Forestry, Sichuan Agricutural University, Yaan, China 2 Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China 3 Department of Biological Sciences, University of Manitoba, Winnipeg, Canada Keywords ABSTRACT Cardiocrinum; floral display; flower number; flower size; trade-off. The evolution of floral display is thought to be constrained by trade-offs between the size and number of flowers; however, empirical evidence for the trade-off is Correspondence inconsistent. We examined evidence for trade-offs and hierarchical allocation of G.-X. Cao, Department of Forestry, Sichuan resources within and between two populations each of the monocarpic perennials, Agricutural University, Yaan 625014, China. Cardiocrinum cordatum and C. giganteum. Within all populations, flower size–num- E-mail: [email protected] ber trade-offs were evident after accounting for variation in plant size. In addition, variation in flower size explained much variation in flower-level allocation to attrac- Editor tion, and female and male function, a pattern consistent with hierarchical alloca- J. Arroyo tion. However, between population differences in flower size (C. cordatum) and number (C. giganteum) were not consistent with size–number trade-offs or hierar- Received: 8 November 2011; Accepted: 19 chical allocation. The population-level difference in C. cordatum likely reflects the March 2012 combined influence of a time lag between initiation and maturation of flowers, and higher light levels in one population. Thus, our study highlights one mechanism doi:10.1111/j.1438-8677.2012.00612.x that may account for the apparent independence of flower size and number in many studies. A prediction of sex allocation theory was also supported. In C. giganteum: plants from one population invested more mass in pistils and less in stamens than did plants from the other population. Detection of floral trade-offs in Cardiocrinum may be facilitated by monocarpic reproduction, production of a single inflorescence and ease of measuring plant size. ume and number per flower (Vonhof & Harder 1995; Worley INTRODUCTION & Barrett 2000; Sarkissian & Harder 2001; Yang & Guo Natural selection favours the allocation of finite resources to 2004) and seed size and number (Leishman 2001; Roff 2002). different functions in a way that maximises plant fitness By contrast, empirical evidence for trade-offs between flower (Charnov 1982). For a given amount of resources, increased size and number is mixed, with reported relationships rang- allocation to one function should necessitate a decrease to ing from positive to non-significant, to negative (Morgan another, thereby resulting in negative relations or trade-offs 1998; Burd 1999; Worley & Barrett 2000; Worley et al. 2000; between competing plant parts (Lloyd 1987; Roff 2002). Caruso 2004; Delph et al. 2004; Tomimatsu & Ohara 2006; Trade-offs that are widely assumed in theoretical treatments Sargent et al. 2007; Goodwillie et al. 2010). For example, Sar- of plant resource allocation include trade-offs between the gent et al. (2007) found a significant negative correlation size and number of repeated organs such as flowers, seeds between flower size and daily flower number in an indepen- and pollen grains (Cohen & Dukas 1990; Morgan 1993; dent contrasts analysis involving 251 species, but did not Harder & Barrett 1995; Sakai 1995, 2000; Schoen & Ashman detect such trade-offs within the genera Collinsia and Narcis- 1995), between female and male function (Parachinowitsch & sus. In some cases the sign and significance of the correlation Elle 2004; Ashman & Majetic 2006) and between current and between flower size and number varies with population and future reproduction (Charnov 1982; Charlesworth & Morgan plant age (e.g. Worley & Barrett 2001; Caruso et al. 2012). 1991). In animal-pollinated species, resource allocation at Empirical evidence is also mixed for trade-offs between flowering should depend on how flower size and number female and male function, especially at flowering (Fenster & affect pollinator attraction, pollination efficiency and the Carr 1997; Parachinowitsch & Elle 2004; Ashman & Majetic associated marginal fitness gains through female and male 2006). function, but trade-offs are expected to place an important Positive or a lack of a correlation between floral traits in constraint on the trait combinations that evolve. many studies suggests that the importance of trade-offs in The empirical evidence suggests that some trade-offs are theoretical models may be overstated. However, multiple sit- indeed ubiquitous whereas evidence for other trade-offs is uations may obscure trade-offs. (i) Within-season allocation much more inconsistent. Examples of widespread trade-offs to flower size and number may not be made from a common involving size and number include those between pollen vol- resource pool at a single point in time (Bazzaz et al. 1979). 158 Plant Biology 15 (2013) 158–165 ª 2012 German Botanical Society and The Royal Botanical Society of the Netherlands Cao & Worley Floral allocation in two moncarpic lilies (ii) Flower size–number trade-offs may only be evident when top to bottom of the inflorescence (basipetal blooming) in lifetime flower production is considered (Morgan 1998; Wor- C. giganteum, whereas almost all flowers within inflorescences ley & Barrett 2000; but see Caruso et al. 2012). (iii) Flower- open simultaneously in C. cordatum (Cao & Kudo 2008). ing and fruiting may draw resources from the same resource Both species are self compatible, but are visited by a wide pool so that negative relations between size and number may variety of insects (Ohara et al. 2006; Cao, unpublished obser- only be apparent if seed production is included as a compo- vations). Fruits mature at the end of September; each fruit nent of investment per flower (Sato & Yahara 1999). (iv) contains several hundred seeds with thin filmy wings in both Variation in plant size (overall resource availability) or hier- species. archical allocation may mask trade-offs between the alloca- tion of resources to flower size and number, or to female Data collection and male function (Van Noordwijk & de Jong 1986; Houle 1991; de Laguerie et al. 1991; de Jong 1993; Koelewijn & We sampled 30 plants selected to span the full size range of Hunscheid 2000; Worley et al. 2000, 2003; Caruso et al. flowering plants in each of two C. cordatum populations on 2012). For example, higher allocation to flowering may allow the Campus of Hokkaido University (43°05¢ N, 141°20¢ E) in outcrossing taxa to produce both more and larger flowers, Sapporo, northern Japan in July 2005. One population was and also to invest more floral resources in both female and located in the understorey of a developed deciduous forest male function than do selfing taxa (Ashman & Majetic 2006; (hereafter Cpop-1). The other population (hereafter Cpop-2) Tomimatsu & Ohara 2006; Goodwillie et al. 2010). was located in a relatively open site with sparse canopy cover. In this study, we eliminated or examined three of the four Relative light intensity was 9.7% in Cpop-1 and 44.3% in situations referred to above. We studied two populations Cpop-2. When flowers began to open, three flowers repre- each of the monocarpic perennials, Cardiocrinum cordatum senting each relative position (basal, middle and distal) (Thunb.) Makino and C. giganteum (Wall.) Makino. Mono- within inflorescences were collected. Flowers were separated carpy, production of a single inflorescence and large plant into stamens, pistils and tepals. The number of flowers was size allowed us to accurately estimate lifetime allocation to counted, and plant size was estimated as a product of square flowering in a single season (situation 2). However, differen- stem diameter · stem height (d2 · h, cm3), which was highly 2 tiation of flowers in Cardiocrinum occurs in autumn so that correlated with total vegetative mass (R = 0.94, F1,19 = flower number is determined several months before flowers 298.91, P < 0.000). All reproductive components were oven are fully mature; this raises the possibility that flower size dried to a constant mass and weighed. The data on C. corda- and number depend on different resource pools (situation 1). tum have been used in a previous study (Cao & Kudo 2008), We compared two populations of C. cordatum that differed but allocation to floral display and evidence for hierarchical markedly in light availability and therefore the resources allocation were not examined. available to developing flowers. Finally, measuring plant size We sampled 29 and 28 plants of C. giganteum encompass- and within-flower allocation allowed us to assess whether ing the full size range of flowering plants in BiFengXia allocation patterns were consistent with a two-level allocation (30°04¢ N, 102°59¢ E) and WangYu (29°45¢ N, 102°56¢ E) hierarchy, in which resources were allocated first between (hereafter Gpop-1 and Gpop-2, respectively), YaAn District, flower size and number and then among attractive, female Sichuan Province, China, in mid-May 2006. All flowers and male functions within flowers (situation 4). Replicate within inflorescences on each plant were collected as they species and populations allowed us to examine the consis- opened. The inflorescence was divided into basal, middle and tency of allocation patterns within this genus. distal portions. All flowers were separated into stamens, pis- We tested the following specific predictions. (i) Flower size tils and tepals, and the average per flower mass of each com- and number both depend on the resources available for flow- ponent was estimated for each position. At the same time, ering, as indicated by plant size. (ii) For a given plant size, the flower number of each individual plant was recorded. All increases in flower size should be at the expense of flower above- and belowground vegetative parts were collected, oven number because these two components of floral allocation dried to a constant mass and weighed.
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