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Nature 515, 108–111 (6 November 2014) | doi:10.1038/nature13869
http://www.natureasia.com/ja-jp/nature/highlights/57462
Nature 515, 7525
2014年11月6日
植物群落は、多様性が高いほど安定性や生産性が高い場合が多く、この効果は時間の経過とともに強まる傾向がある。今回D Zuppinger-Dingleyたちは、この生態学的現象に進化が果たす役割を検討した。それぞれ混合栽培と単独栽培とで育てた、最近の進化的履歴が異なる植物(イネ科植物、双子葉草本類およびマメ科植物)を組み合わせて、新たな生態学的群落を構築した。生態系機能および形態形質の多様性は、混合状態で栽培された植物の群落の方が単独で栽培された植物の群落よりも高かったことから、個体群が生物的環境に適応していくにつれて自然群落の生物多様性効果が強まることが示唆された。小規模な進化が生態的関係に重要であるという今回の知見は、自然群落中の種の共存に対するこうしたさまざまな観点を総合するものである。
Letter p.108
Selection for niche differentiation in plant communities increases biodiversity effects
Debra Zuppinger-Dingley, Bernhard Schmid, Jana S. Petermann, Varuna Yadav, Gerlinde B. De Deyn
& Dan F. B. Flynn
Affiliations
Contributions
Corresponding authors
Nature
515,
108–111
(06 November 2014)
doi:10.1038/nature13869
Received 11 June 2014
Accepted 17 September 2014
Published online 15 October 2014
In experimental plant communities, relationships between biodiversity and ecosystem functioning have been found to strengthen over time1, 2, a fact often attributed to increased resource complementarity between species in mixtures3 and negative plant–soil feedbacks in monocultures4. Here we show that selection for niche differentiation between species can drive this increasing biodiversity effect. Growing 12 grassland species in test monocultures and mixtures, we found character displacement between species and increased biodiversity effects when plants had been selected over 8 years in species mixtures rather than in monocultures. When grown in mixtures, relative differences in height and specific leaf area between plant species selected in mixtures (mixture types) were greater than between species selected in monocultures (monoculture types). Furthermore, net biodiversity and complementarity effects1, 2 were greater in mixtures of mixture types than in mixtures of monoculture types. Our study demonstrates a novel mechanism for the increase in biodiversity effects: selection for increased niche differentiation through character displacement. Selection in diverse mixtures may therefore increase species coexistence and ecosystem functioning in natural communities and may also allow increased mixture yields in agriculture or forestry. However, loss of biodiversity and prolonged selection of crops in monoculture may compromise this potential for selection in the longer term.
Figure 1: Experimental design.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_F1.html
From
Selection for niche differentiation in plant communities increases biodiversity effects

Plant material, shoots and roots (n = 4,900) from experimental field monoculture versus mixture selection communities established in Jena, Germany in 2002, were collected in 2010 and assembled in new experimental glasshouse monoculture versus mixture test communities in 2010/11 (n = 855). We expected that mixture test communities would have higher productivity if assembled from plants collected from mixture selection communities in the field and vice versa for monocultures. Different shades of green represent the hypothesized selection for monoculture or mixture types from 2002 to 2010.
Figure 2: Stronger biodiversity effects for plants selected in mixtures compared with plants selected in monocultures.
From Selection for niche differentiation in plant communities increases biodiversity effects

a, b, Plants selected in mixture plots in the Jena Experiment over 8 years showed stronger biodiversity effects than plants selected in monoculture plots over the same time period. Mix, mixture types; Mono, monoculture types. a, Biodiversity effects were assessed by additive partitioning15 of net effects into complementarity and sampling effects (n = 545). b, The plots are ordered by functional group combinations: grasses (g); small herbs (sh); tall herbs (th); legumes (l). Symbols are means ±1 standard error of the mean (s.e.m.) calculated from raw data.
Figure 3: Plants selected in mixtures show character displacement between species when grown in mixture.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_F3.html
From Selection for niche differentiation in plant communities increases biodiversity effects

a–c, After 8 years of selection in the Jena Experiment, mixture types (Mix) in comparison with monoculture types (Mono) showed character displacement between species. a, b, Relative differences between the two species in mixture for plant height (n = 219 aggregated differences) (a) and SLA (n = 208 aggregated differences) (b). c, Functional diversity30 (calculated from height, SLA and reproductive biomass) of the two species in mixture (unit-less functional diversity index, n = 219 aggregated values). Symbols are means ±1 s.e.m. calculated from raw data. Means are averages over all 50 species combinations.
Extended Data Figure 1: Designed number of pots planted for each combination of species.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_SF1.html
From Selection for niche differentiation in plant communities increases biodiversity effects

Plants from three different selection histories (monoculture, plot containing one species; monofunctional group, plot containing at least four species of the same functional group of plants; mixed-functional group, plot containing at least four species of four different functional groups) were grown in three different types of test communities (monoculture, monofunctional mixture, mixed-functional mixture). Twelve species in the Jena Experiment were chosen from all four functional groups: grass (g) (Festuca pratensis, Festuca rubra, Poa pratensis), small herb (sh) (Plantago lanceolata, Prunella vulgaris, Veronica chamaedrys), tall herb (th) (Crepis biennis, Galium mollugo, Geranium pratense), legume (l) (Lathyrus pratensis, Onobrychis viciifolia, Trifolium repens); numbers after the letter abbreviations refer to the different species. This design was used once with plants raised from cuttings (Block 1) and once with plants raised from seedlings (Block 2). Overall we aimed to obtain the same 12 monocultures and 48 two-species combinations as test communities for each block. Availability of species precluded some of the two-species combinations in each block, such that they had to be replaced by other combinations. This yielded a total of 50 combinations across the two blocks, with several that were unique within a block. Each monoculture and each two-species combination was assembled three times for each of the three types of selection histories in each block. Some monocultures and some two-species combinations could not be realized with all types of selection histories in both blocks. Overall, there were 855 pots, 168 monocultures and 687 two-species mixtures; for 545 of the latter, the net biodiversity effect could be partitioned into complementarity and sampling effects. Some missing monocultures precluded the calculation of biodiversity effects in certain mixtures.
Extended Data Figure 2: Biodiversity–productivity relationship is stronger for plants with a common selection history.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_SF2.html
From Selection for niche differentiation in plant communities increases biodiversity effects

Aboveground net primary productivity of communities in an experimental manipulation of plant species richness and selection history (common history versus no common history). In this experiment, species represented an expanded set from the present experiment (52 species), and were planted within a large-scale field experiment in Jena, Germany, on mixed soil from 48 plots from which plants had been selected, thus equalizing potential effects of soil legacy among treatments. Plants without selection history were grown from seed from a seed company, while plants with selection history were seed progeny from plots of exactly the same species composition as the one in which they were replanted (same propagation procedure as for the 12 species used in the test communities of the present study). The slope of the biodiversity–productivity relationship was steeper for plants with a common selection history (significance of slope differences tested with interaction term log(species richness) × selection history in mixed model with random-effects factor for 48 specific plant communities; P < 0.001, n = 96).
Extended Data Figure 3: Selection for different biochemical features in monocultures and mixtures.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_SF3.html
From
Selection for niche differentiation in plant communities increases biodiversity effects
Debra Zuppinger-Dingley, Bernhard Schmid, Jana S. Petermann, Varuna Yadav, Gerlinde B. De Deyn & Dan F. B. Flynn
Nature 515, 108–111 (06 November 2014) doi:10.1038/nature13869

Ordinations (non-metric multidimensional scaling (NMDS)) of second derivative of spectral wavenumbers of 8 of the 12 species used in the present study, showing effects of 8-year selection history on plant individuals derived from monoculture and mixture communities (Jena Experiment). This can be an indication of selection for different biochemical features over 8 years in monoculture and mixtures. Stress values reflect a measure of goodness of fit for NMDS, with lower values showing better representation of the original data.
Nature 515, 108–111 (6 November 2014) | doi:10.1038/nature13869
http://www.natureasia.com/ja-jp/nature/highlights/57462
Nature 515, 7525
2014年11月6日
植物群落は、多様性が高いほど安定性や生産性が高い場合が多く、この効果は時間の経過とともに強まる傾向がある。今回D Zuppinger-Dingleyたちは、この生態学的現象に進化が果たす役割を検討した。それぞれ混合栽培と単独栽培とで育てた、最近の進化的履歴が異なる植物(イネ科植物、双子葉草本類およびマメ科植物)を組み合わせて、新たな生態学的群落を構築した。生態系機能および形態形質の多様性は、混合状態で栽培された植物の群落の方が単独で栽培された植物の群落よりも高かったことから、個体群が生物的環境に適応していくにつれて自然群落の生物多様性効果が強まることが示唆された。小規模な進化が生態的関係に重要であるという今回の知見は、自然群落中の種の共存に対するこうしたさまざまな観点を総合するものである。
Letter p.108
Selection for niche differentiation in plant communities increases biodiversity effects
Debra Zuppinger-Dingley, Bernhard Schmid, Jana S. Petermann, Varuna Yadav, Gerlinde B. De Deyn
& Dan F. B. Flynn
Affiliations
Contributions
Corresponding authors
Nature
515,
108–111
(06 November 2014)
doi:10.1038/nature13869
Received 11 June 2014
Accepted 17 September 2014
Published online 15 October 2014
In experimental plant communities, relationships between biodiversity and ecosystem functioning have been found to strengthen over time1, 2, a fact often attributed to increased resource complementarity between species in mixtures3 and negative plant–soil feedbacks in monocultures4. Here we show that selection for niche differentiation between species can drive this increasing biodiversity effect. Growing 12 grassland species in test monocultures and mixtures, we found character displacement between species and increased biodiversity effects when plants had been selected over 8 years in species mixtures rather than in monocultures. When grown in mixtures, relative differences in height and specific leaf area between plant species selected in mixtures (mixture types) were greater than between species selected in monocultures (monoculture types). Furthermore, net biodiversity and complementarity effects1, 2 were greater in mixtures of mixture types than in mixtures of monoculture types. Our study demonstrates a novel mechanism for the increase in biodiversity effects: selection for increased niche differentiation through character displacement. Selection in diverse mixtures may therefore increase species coexistence and ecosystem functioning in natural communities and may also allow increased mixture yields in agriculture or forestry. However, loss of biodiversity and prolonged selection of crops in monoculture may compromise this potential for selection in the longer term.
Figure 1: Experimental design.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_F1.html
From
Selection for niche differentiation in plant communities increases biodiversity effects

Plant material, shoots and roots (n = 4,900) from experimental field monoculture versus mixture selection communities established in Jena, Germany in 2002, were collected in 2010 and assembled in new experimental glasshouse monoculture versus mixture test communities in 2010/11 (n = 855). We expected that mixture test communities would have higher productivity if assembled from plants collected from mixture selection communities in the field and vice versa for monocultures. Different shades of green represent the hypothesized selection for monoculture or mixture types from 2002 to 2010.
Figure 2: Stronger biodiversity effects for plants selected in mixtures compared with plants selected in monocultures.
From Selection for niche differentiation in plant communities increases biodiversity effects

a, b, Plants selected in mixture plots in the Jena Experiment over 8 years showed stronger biodiversity effects than plants selected in monoculture plots over the same time period. Mix, mixture types; Mono, monoculture types. a, Biodiversity effects were assessed by additive partitioning15 of net effects into complementarity and sampling effects (n = 545). b, The plots are ordered by functional group combinations: grasses (g); small herbs (sh); tall herbs (th); legumes (l). Symbols are means ±1 standard error of the mean (s.e.m.) calculated from raw data.
Figure 3: Plants selected in mixtures show character displacement between species when grown in mixture.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_F3.html
From Selection for niche differentiation in plant communities increases biodiversity effects

a–c, After 8 years of selection in the Jena Experiment, mixture types (Mix) in comparison with monoculture types (Mono) showed character displacement between species. a, b, Relative differences between the two species in mixture for plant height (n = 219 aggregated differences) (a) and SLA (n = 208 aggregated differences) (b). c, Functional diversity30 (calculated from height, SLA and reproductive biomass) of the two species in mixture (unit-less functional diversity index, n = 219 aggregated values). Symbols are means ±1 s.e.m. calculated from raw data. Means are averages over all 50 species combinations.
Extended Data Figure 1: Designed number of pots planted for each combination of species.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_SF1.html
From Selection for niche differentiation in plant communities increases biodiversity effects

Plants from three different selection histories (monoculture, plot containing one species; monofunctional group, plot containing at least four species of the same functional group of plants; mixed-functional group, plot containing at least four species of four different functional groups) were grown in three different types of test communities (monoculture, monofunctional mixture, mixed-functional mixture). Twelve species in the Jena Experiment were chosen from all four functional groups: grass (g) (Festuca pratensis, Festuca rubra, Poa pratensis), small herb (sh) (Plantago lanceolata, Prunella vulgaris, Veronica chamaedrys), tall herb (th) (Crepis biennis, Galium mollugo, Geranium pratense), legume (l) (Lathyrus pratensis, Onobrychis viciifolia, Trifolium repens); numbers after the letter abbreviations refer to the different species. This design was used once with plants raised from cuttings (Block 1) and once with plants raised from seedlings (Block 2). Overall we aimed to obtain the same 12 monocultures and 48 two-species combinations as test communities for each block. Availability of species precluded some of the two-species combinations in each block, such that they had to be replaced by other combinations. This yielded a total of 50 combinations across the two blocks, with several that were unique within a block. Each monoculture and each two-species combination was assembled three times for each of the three types of selection histories in each block. Some monocultures and some two-species combinations could not be realized with all types of selection histories in both blocks. Overall, there were 855 pots, 168 monocultures and 687 two-species mixtures; for 545 of the latter, the net biodiversity effect could be partitioned into complementarity and sampling effects. Some missing monocultures precluded the calculation of biodiversity effects in certain mixtures.
Extended Data Figure 2: Biodiversity–productivity relationship is stronger for plants with a common selection history.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_SF2.html
From Selection for niche differentiation in plant communities increases biodiversity effects

Aboveground net primary productivity of communities in an experimental manipulation of plant species richness and selection history (common history versus no common history). In this experiment, species represented an expanded set from the present experiment (52 species), and were planted within a large-scale field experiment in Jena, Germany, on mixed soil from 48 plots from which plants had been selected, thus equalizing potential effects of soil legacy among treatments. Plants without selection history were grown from seed from a seed company, while plants with selection history were seed progeny from plots of exactly the same species composition as the one in which they were replanted (same propagation procedure as for the 12 species used in the test communities of the present study). The slope of the biodiversity–productivity relationship was steeper for plants with a common selection history (significance of slope differences tested with interaction term log(species richness) × selection history in mixed model with random-effects factor for 48 specific plant communities; P < 0.001, n = 96).
Extended Data Figure 3: Selection for different biochemical features in monocultures and mixtures.
http://www.nature.com/nature/journal/v515/n7525/fig_tab/nature13869_SF3.html
From
Selection for niche differentiation in plant communities increases biodiversity effects
Debra Zuppinger-Dingley, Bernhard Schmid, Jana S. Petermann, Varuna Yadav, Gerlinde B. De Deyn & Dan F. B. Flynn
Nature 515, 108–111 (06 November 2014) doi:10.1038/nature13869

Ordinations (non-metric multidimensional scaling (NMDS)) of second derivative of spectral wavenumbers of 8 of the 12 species used in the present study, showing effects of 8-year selection history on plant individuals derived from monoculture and mixture communities (Jena Experiment). This can be an indication of selection for different biochemical features over 8 years in monoculture and mixtures. Stress values reflect a measure of goodness of fit for NMDS, with lower values showing better representation of the original data.