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The effects of nutrient addition on floral characters and pollination in two subalpine plants, Ipomopsis aggregata and Linum lewisii

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Abstract

The availability of soil and pollination resources are main determinants of fitness in many flowering plants, but the degree to which each is limiting and how they interact to affect plant fitness is unknown for many species. We performed resource (water and nutrients) and pollination (open and supplemental) treatments on two species of flowering plants, Ipomopsis aggregata and Linum lewisii, that differed in life-history, and we measured how resource addition affected floral characters, pollination, and reproduction (both male and female function). We separated the direct effects of resources versus indirect effects on female function via changes in pollination using a factorial experiment and path analysis. Resource addition affected I. aggregata and L. lewisii differently. Ipomopsis aggregata, a monocarp, responded to fertilization in the year of treatment application, increasing flower production, bloom duration, corolla width, nectar production, aboveground biomass, and pollen receipt relative to control plants. Fertilization also increased total seed production per plant, and hand-pollination increased seeds per fruit in I. aggregata, indicating some degree of pollen limitation of seed production. In contrast, fertilization had no effect on growth or reproductive output in the year of treatment on L. lewisii, a perennial, except that fertilization lengthened bloom duration. However, delayed effects of fertilization were seen in the year following treatment, with fertilized plants having greater aboveground biomass, seeds per fruit, and seeds per plant than control plants. In both species, there were no effects of resource addition on male function, and the direct effects of fertilization on female function were relatively stronger than the indirect effects via changes in pollination. Although we studied only two plant species, our results suggest that life-history traits may play an important role in determining the reproductive responses of plants to soil nutrient and pollen additions.

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Acknowledgments

We thank P. McDonald for field assistance and the Irwin lab group for discussion and support. W. Bowman, S. Elliott, M. McPeek, P. Muller, M. Price, L. Rolfe, and three anonymous reviewers provided insightful comments on earlier drafts of this manuscript. Field research was funded by a grant from the Colorado Mountain Club, and supplies were provided by a grant from the National Science Foundation (DEB-0089643).

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  1. Department of Biological Sciences, Dartmouth College, Hanover, NH, 03755, USA

    Laura A. Burkle & Rebecca E. Irwin

  2. Rocky Mountain Biological Laboratory, Crested Butte, CO, 81224, USA

    Laura A. Burkle & Rebecca E. Irwin

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  1. Laura A. Burkle
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Correspondence to Laura A. Burkle.

Appendix 1. Comparison of effect sizes for Ipomopsis aggregata between 1990 and 2004

Appendix 1. Comparison of effect sizes for Ipomopsis aggregata between 1990 and 2004

In 1990, Campbell and Halama (1993) manipulated soil resources and pollination to I. aggregata at the RMBL using the same nutrient addition amounts and schedules and measured similar response variables as were measured in this study. Thus, we were provided with an opportunity to test whether the effects of resources and pollen were consistent across years at the exact same study site at the RMBL. We calculated and compared effect sizes of I. aggregata in the two studies using the standardized mean difference, d (Hedges et al. 1999), the difference between the means of two treatments divided by their pooled standard deviation. A positive effect size indicated positive effects of the treatments and vice versa. An effect size of 0.2 was considered small, 0.5 medium, and greater than 0.8 large (Cohen 1969). Because the water treatment had minimal effects on floral characters and female plant reproduction in Campbell and Halama (1993) and in this study (see “Results”), we limited our comparison to fertilized and control treatments. For corolla width and nectar production, we compared the fertilized and control treatments of each of the two studies. For flower production, seeds per fruit, and total seeds per plant, we compared all factorial combinations of nutrient and pollination treatments.

Overall, we found similar directional responses to nutrient and pollination treatments between the 1990 (Campbell and Halama 1993) and 2004 (here) studies, although some effect sizes varied in magnitude (Table A). Large effect sizes (d > 0.8) of nutrient and/or pollen treatments were calculated for most comparisons. Fertilization in this study resulted in larger effect sizes on some floral traits than in Campbell and Halama (1993). In particular, the effect size for corolla width was 5 times larger and for nectar production 4.4 times larger in 2004 than 1990. There were only two comparisons, between total seeds of fertilized open- versus hand-pollinated plants and seeds per fruit of open-pollinated fertilized versus control plants, in which we saw an effect in this study where none was observed in 1990. In addition, there was only one comparison, between total flowers of fertilized open- versus hand-pollinated plants, in which the direction of effect changed between studies (negative in 1990 to positive in 2004). These two studies indicate that nutrients and pollen are limiting resources for I. aggregata in two different years, but the magnitude of limitation was generally greater in 2004 than in 1990.

Table A Effect sizes of nutrient and pollination treatments in 1990 (Campbell and Halama 1993) and 2004 (this study)
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Burkle, L.A., Irwin, R.E. The effects of nutrient addition on floral characters and pollination in two subalpine plants, Ipomopsis aggregata and Linum lewisii . Plant Ecol 203, 83–98 (2009). https://doi.org/10.1007/s11258-008-9512-0

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