Abstract
When plant monocultures are sown over a wide range of densities for a given period of time, the total biomass yield increases with density at low densities and then levels off at high densities, a phenomenon called constant final yield (CFY). There are several reported cases, however, where the total yield decreases at very high densities, but the reasons for such exceptions are not known. We used a spatially explicit, individual-based “field of neighborhood” simulation model to investigate the potential roles of spatial pattern, individual variation, and competitive stress tolerance for CFY. In the model, individual plants compete asymmetrically for light when their fields overlap, and this competition decreases growth and increases mortality. We varied (1) the initial size variation, (2) the spatial pattern, and (3) ability to survive intense competition and examined the effects on the density-biomass relationship. CFY was always observed when there was high variability among individuals, but not always when variability was low. This high size variation could be the result of high initial size variability or variation in the degree of local crowding. For very different reasons, very high and very low tolerance for competition resulted in decreasing total biomass at very high densities. Our results emphasize the importance of individual variation for population processes and suggest that we should look for exceptions to CFY in homogeneous, even-aged, regularly spaced populations such as plantations.
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References
Bagchi R, Swinfield T, Gallery RE, Lewis OT, Gipenger S, Narayan L, Freckleton RP (2010) Testing the Janzen-Connell mechanism: pathogens causes overcompensating density dependence in the a tropical tree. Ecol Lett 13:1262–1269
Bazzaz FA, Harper JL (1976) Relationship between plant weight and numbers in mixed populations of Sinapis alba (L.) Rabenh. and Lepidium sativum L. J Appl Ecol 13:211–216
Berger U, Hildenbrandt H (2000) A new approach to spatially explicit modelling of forest dynamics: spacing, ageing and neighbourhood competition of mangrove trees. Ecol Model 132:287–302
Berger U, Hildenbrandt H, Grimm V (2004) Age-related decline in forest production: modelling the effects of growth limitation, neighbourhood competition and self-thinning. J Ecol 92:846–853
Berger U, Adams M, Grimm V, Hildenbrandt H (2006) Modelling secondary succession of neotropical mangroves: causes and consequences of growth reduction in pioneer species. Perspect Plant Ecol Evol Syst 7:243–252
Bleasdale JKA (1966) The effects of plant spacing on the yield of bulb onion (Allium cepa L.) grown from seed. J Hortic Sci 41:145–153
Bonan GB (1991) Density effects on the size structure of annual plant populations: an indication of neighbourhood competition. Ann Bot 68:341–347
Chu CJ, Maestre FT, Xiao S, Weiner J, Wang YS, Duan ZH, Wang G (2008) Biomass-density relationships in plant populations are determined by the balance between facilitation and resource competition. Ecol Lett 11:1189–1197
Chu CJ, Weiner J, Maestre FT, Xiao S, Wang YS, Li Q, Yuan JL, Zhao LQ, Ren ZW, Wang G (2009) Positive interactions can increase size inequality in plant populations. J Ecol 97:1401–1407
Farazdaghi H, Harris PM (1968) Plant competition and crop yield. Nature 217:289–290
Fibich P, Lepš (2011) Do biodiversity indices behave as expected from traits of constituent species in simulated scenarios? Ecol Model 222:2049–2058
Grimm V, Railsback SF (2005) Individual-based modeling and ecology. University Press, Princeton
Hara T, Wyszomirski T (1994) Competition asymmetry reduces spatial effects on size-structure dynamics in plant populations. Ann Bot 73:285–297
Kays S, Harper JL (1974) The regulation of plant and tiller density in a grass sward. J Ecol 62:97–105
Kotorova I, Leps J (1999) Comparative ecology of seedling recruitment in an oligotrophic wet meadow. J Veg Sci 10:175–186
Kristensen L, Olsen J, Weiner J (2008) Crop density, sowing pattern and nitrogen fertilization effects on weed suppression and yield in spring wheat. Weed Sci 56:97–102
Lepš J, Kindlmann P (1987) Models of development of spatial pattern of an even-aged plant population over time. Ecol Model 39:45–89
Li B, Watkinson AR, Hara T (1996) Dynamics of competition in populations of carrot (Daucus carota). Ann Bot 78:203–214
Miller TE, Weiner J (1989) Local density variation may mimic effects of asymmetric competition on plant size variability. Ecology 70:1188–1191
Pacala SA, Weiner J (1991) Effects of including competitive asymmetry in a local density model of plant interference. J Theor Biol 149:165–179
Prach K (1982) Self-thinning processes in selected ruderal species populations. Preslia 54:271–275
Pretzsch H (2003) The elasticity of growth in pure and mixed stands of Norway spruce (Picea abies L. Karst.) and common beech (Fagus sylvatica L.). J For Sci 49:491–501
Rahman MM, Hossain MM (2011) Plant density effects on growth, yield and yield components of two soybean varieties under equidistant planting arrangement. Asian J Plant Sci 10:278–286
Reynolds JD (1950) Spacing trials with dried peas. Agriculture 56:527–537
Rumpel J, Felczynski K (2000) Effect of plant density on yield and bulb size of direct sown onions. Acta Horticult 533:179–186
Scaife MA, Jones D (1976) The relationship between crop yield (or mean plant weight) of lettuce and plant density, length of growing period, and initial plant weight. J Agric Sci 86:83–91
Shinozaki K, Kira T (1956) Intraspecific competition among higher plants. VII. Logistic theory of the C-D effect. J Inst Polytechnics Osaka City Univ 7:35–72
Silvertown J, Charlesworth D (2001) Introduction to plant population biology. Blackwell, London
Stachová T, Fibich P, Lepš J (2013) Plant density affects measures of biodiversity effects. J Plant Ecol 6:1–11
Stoker R (1975) Effect of plant population on yield garden peas under different moisture regimes. N Z J Exp Agric 3:333–337
Stoll P, Weiner J, Muller-Landau H, Müller E, Hara T (2002) Size symmetry of competition alters biomass-density relations. Proc R Soc B 296:2191–2195
Symonides E (1983a) Population size regulation as a result of intra-population interactions 1. Effect of density on the survival and development of individuals of Erophila verna (L.). Ekologia Polska 31:839-882
Symonides E (1983b) Population size regulation as a result of intra-population interactions 2. Effect of density on the growth rate morphological diversity and fecundity of Erophila verna (L.). Ekologia Polska 31:883-912
Taylor DR, Aarssen LW (1989) On the density dependence of replacement-series competition experiments. J Ecol 4:975–988
Uchmanski J (2000) Individual variability and population regulation: an individual-based model. Oikos 90:539–548
Weiner J, Freckleton R (2010) Constant final yield. Annu Rev Ecol Evol Syst 41:173–192
Weiner J, Thomas SC (1986) Size variability and competition in plant monocultures. Oikos 47:211–222
Weiner J, Thomas SC (1992) Competition and allometry in three species of annual plants. Ecology 73:648–656
Weiner J, Kinsman S, Williams S (1998) Modeling the growth of individuals in plant populations: local density variation in a strand population of Xanthium strumarium. Am J Bot 85:1638–1645
Weiner J, Stoll P, Muller-Landau H, Jasentuliyana A (2001) The effects of density, spatial pattern and competitive symmetry on size variation in simulated plant populations. Am Nat 158:438–450
Willey RW, Heath SB (1969) The quantitative relationships between plant population and crop yield. Adv Agron 24:281–322
Xue L, Hagihara A (2008) Density effects on organs in self-thinning Pinus densiflora Sieb. Et Zucc. stands. Ecol Res 23:689–695
Yahuza I (2011) Yield-density equations and their application for agronomic research: a review. Int J Biosci 1:1–17
Yastrebov AB (1996) Different types of heterogeneity and plant competition in monospecific stands. Oikos 75:89–97
Yoda K, Kira T, Ogawa H, Hozumi K (1963) Self-thinning in overcrowded pure stands under cultivated and natural conditions. J Biol Osaka City Univ 14:107–129
Acknowledgement
The study was supported by the grant GACR 13-17118S from the Grant Agency of the Czech Republic and the University of Copenhagen Program of Excellence. Access to computing and storage facilities owned by parties and projects contributing to the National Grid Infrastructure MetaCentrum provided under the program “Projects of Large Infrastructure for Research, Development, and Innovations” (LM2010005) is much appreciated.
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Fibich, P., Lepš, J. & Weiner, J. Individual variability and mortality required for constant final yield in simulated plant populations. Theor Ecol 7, 263–271 (2014). https://doi.org/10.1007/s12080-014-0216-x
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DOI: https://doi.org/10.1007/s12080-014-0216-x