Effects of Crop Diversity on Agroecosystem Function: Crop Yield Response

Understanding the role of diversity in the functioning of ecosystems has important implications for agriculture. Previous agricultural research has shown that crop rotation and the use of cover crops can lead to increases in yield relative to monoculture; however, few studies have been performed within the broader context of diversity–ecosystem function theory and in the absence of chemical inputs. We performed a field experiment in SW Michigan, USA, in which we manipulated the number of crop species grown in rotation and as winter cover crops over a 3-year period to test if varying the number of species in a rotation affected grain yield, a critical metric of ecosystem function in row-crops. The experimental design was unique in that no fertilizer or pesticides were used, and the only management variable manipulated was number of species in the rotation, thus providing a strong comparison to grassland diversity–ecosystem function experiments. Treatments included continuous monocultures of three row-crops, corn Zea mays L., soybean Glycine max (L.) Merr., and winter wheat Triticum aestivum L., and 2- and 3-year annual rotations with and without cover crops (zero, one, or two legume/small grain species), encompassing a range of crop diversity from one to six species. Crop yields and weed biomass were measured annually for 3 years and plant available soil nitrogen was measured over the course of the growing season in the final year of the study. In all 3 years, corn grain yield increased linearly in response to the number of crops in the rotation. Corn yields in the highest diversity treatment (three crops, plus three cover crops) were over 100% higher than in continuous monoculture and were not significantly different from the county average for each of the 3 years despite the absence of chemical inputs. Corn yields in the diversity treatments were strongly correlated with the availability of inorganic soil nitrogen, which was likely influenced by the number of different legume species (crops and cover crops) present in the rotation. In soybean and winter wheat, yield differences among crop diversity treatments were also significant, but of lower magnitude (32 and 53%, respectively), and showed little direct relationship to the number of crop species grown in a rotation. Results demonstrate that agricultural research motivated by ecological theory can provide important insights into the functioning of agroecosystems and enhance our understating of the linkages between diversity and ecosystem function. Importantly, these results suggest that reduced chemical inputs do not necessarily result in yield penalties and provide support for incorporation of crop or species diversity when determining how ecosystem services can be included in food, fiber, and biofuel production.     

Divergence of mycorrhizal fungal communities in crop production systems

Mycorrhizal fungi are present in all arable soils and colonize nearly all crops and weed pests of crops. They may be involved as mutualists or pathogens of crops in well known but poorly understood phenomena such as crop rotation and green manure effects on soil productivity. Crop change effects on mycorrhizal fungal community parameters were evaluated in three field experiments. In Experiment 1, soybean (Glycine max (L.) Merr. cv. Douglas) was grown continuously or rotated with corn (Zea mays L.), milo (Sorghum bicolor (L.) Moench), or fescue (Festuca arundinacea Schreb cv. Johnstone) for two years, then soybean was grown on all plots. Continuous soybean plots were dominated byGigaspora spp., while rotated crops were dominated byGlomus spp. Differences in communities and community indices of continuous soybean and rotated plots were reduced after growing soybeans on rotated plots. In Experiment 2, a fescue sod was plowed and pearl millet (Pennisetum americanum Leeke) or crabgrass (Digitaria sanguinalis (L.) Scop.) grown. Both hosts resulted in great changes in populations of individual species, decreases in community dominance, and increases in community diversity and equitability. Crabgrass also resulted in reduced species richness. In Experiment 3, tobacco (Nicotiana tabacum L.) or fescue was planted on adjacent tracts of land with a long-term history of either fescue (30 yr) or sorghum-sudangrass (Sorghum bicolor (L.) Moench. ×S. sudanense (Piper) Staph.) (3 yr). The long-term cropping history had major effects on the mycorrhizal fungal communities which were related to the expression of mycorrhizal stunt disease of tobacco. Changes occurred in these communities in response to either current-season crop. These experiments suggest that crop rotation causes large changes in mycorrhizal fungal communities, that these changes may be involved in the rotation effect on soil productivity, and that design of cropping systems should take mycorrhizal fungal communities into consideration.     

不同作物轮作制度对土壤杂草种子库特征的影响

为明确不同作物轮作制度对杂草种子库特征的影响,比较研究了玉米-小麦(MW)、大豆-小麦(SW)轮作和原水稻-小麦(RW)轮作方式下杂草种子库的特征。结果表明,作物轮作显著影响杂草种子库的密度和种类组成,进行玉米-小麦或大豆-小麦轮作2年后,杂草种子库密度分别比稻麦轮作降低27．16％、44．44％。在原水稻-小麦轮作方式下,种子库中主要杂草为陌上菜、异型莎草、菵草、水苋菜、千金子等。玉米-小麦轮作使种子库中马唐、碎米莎草、飘拂草的相对优势度显著上升,鸭舌草、水苋菜、菵草的相对优势度显著下降。而在大豆-小麦轮作方式下,种子库中通泉草、马唐、鳢肠、飘拂草的相对优势度显著上升。玉米-小麦轮作种子库的物种多样性指数高于大豆-小麦轮作和水稻-小麦轮作。玉米-小麦与大豆-小麦轮作种子库的物种组成相似性较高。作物轮作影响杂草种子库密度和种类组成的机制,可能在于通过轮换种植不同的作物,提供了多样化的选择压,限制了某些对单一种植系统有着良好适应性的杂草种类的生长。     

Impact of crop management on intraspecific diversity of Pseudomonas corrugata in bulk soil

Colonization at sugar beet root surfaces by seedling-inoculated biocontrol strain Pseudomonas fluorescens DR54 and native soil bacteria was followed over a period of 3 weeks using a combination of immunofluorescence (DR54-targeting specific antibody) and fluorescence in situ hybridization (rRNA-targeting Eubacteria EUB338 probe) techniques with confocal laser scanning microscopy. The dual staining protocol allowed cellular activity (ribosomal number) to be recorded in both single cells and microcolonies of strain DR54 during establishment on the root. After 2 days, the population density of strain DR54 reached a constant level at the root basis. From this time, however, high cellular activity was only found in few bacteria located as single cells, whereas all microcolony-forming cells occurring in aggregates were still active. In contrast, a low density of strain DR54 was observed at the root tip, but here many of the bacteria located as single cells were active. The native population of soil bacteria, comprising a diverse assembly of morphologically different forms and size classes, initiated colonization at the root basis only after 2 days of incubation. Hence the dual staining protocol allowed direct microscopic studies of early root colonization by both inoculant and native soil bacteria, including their differentiation into active and non-active cells and into single or microcolony-forming cells.     

Diversity and phenology of weed communities in the Nile Delta region

This study evaluates the species diversity and phe-nological behaviour of the weed communities associated with the common crops in the Nile Delta region (three winter crops: Egyptian clover, broad beans and wheat; and three summer crops: cotton, maize and rice). The weed communities of winter crops are of higher diversity than those of summer crops, presumably due to differences in sowing and weeding processes. The phenological sequences of the weed communities are, in most cases, closely related to those of the crops in which they occur. The correspondence between the crucial periods in the life cycle of the weed and the crop (seedling establishment and fruiting) often leads to severe competition.     

Competition at the population level along a standing crop gradient: a field experiment in successional grassland

I measured competitive responses of experimentally-established populations of the perennial grass, Andropogon gerardi, across a complex gradient of standing crop and species composition in the successional grasslands of southwest Michigan. The goal was to assess whether long-term (three year) population-level responses of Andropogon to competition matched the inferences made from a previous phytometer study that examined transplant responses to competition across this same gradient over a single growing season.Replicate experimental populations of Andropogon were established at seven grassland sites by sowing seed into 0.5×0.5 m plots that had been denuded of all vegetation. During the first year of the study, all Andropogon populations were maintained as monocultures by hand weeding. At the end of the first growing season, half of the monocultures were selected for continued weeding and half were left open to invasion by competitors for three years. Invasion of the unweeded populations by neighboring plants varied strongly among sites and was positively correlated with standing crop. Increased susceptibility to invasion and competition resulted in the extinction of the unweeded Andropogon populations at the two most productive sites, supporting the hypothesis that Andropogon is restricted by competition to low productivity sites in these grasslands. The finding that the intensity of competition was positively correlated with standing crop is consistent with the previous transplant study, suggesting that short-term experimental assays of competition on the growth of individual transplants may have predictive value for longer-term outcomes of competition at the population level.     

Effects of plant diversity on invasion of weed species in experimental pasture communities

Studies have shown that weed invasion into grasslands may be suppressed if the resident plant community is sufficiently diverse. The objective of this study was to determine whether increased forage plant diversity in grazed pasture communities might be associated with reduced weed abundance both in the aboveground vegetation and soil seed bank. Data were collected from a pasture experiment established in 1994 in Missouri, USA. The experiment consisted of 15 m×15 m plots sown with Festuca arundinacea Schreb. or Bromus inermis Leysser as a base species in mixtures of 1, 2, 3, 6, or 8 forage species. The plots were grazed by cattle during each growing season from 1998 to 2002. Aboveground plant species composition in each plot was measured using a point step method. Soil cores were collected in 1999 and 2002, and the species composition of germinable weed seeds in plots were evaluated by identifying seedlings as they germinated over an 8-week period. Species diversity was measured using several indices: species richness (S), Shannon–Wiener diversity index (H^′), and forage species evenness (J). Aboveground weed abundance in plots was unrelated to forage species richness (S), but weed abundance declined as the evenness (J) of resident forage species increased in mixtures. The species composition of mixtures may have affected weed abundance. Weeds both in the soil seed bank and aboveground vegetation were less abundant in mixtures that contained F. arundinacea compared with mixtures that contained B. inermis. Although variables like forage plant productivity may also suppress weed abundance in pastures, our results suggest that maintaining an evenly distributed mixture of forage species may help suppress weeds as well.

Zusammenfassung

Untersuchungen haben gezeigt, dass die Unkrautinvasion in Grünländer unterdrückt sein kann, wenn die ansässige Pflanzengemeinschaft ausreichend divers ist. Die Zielsetzung dieser Untersuchung war es zu bestimmen, ob eine erhöhte Futterpflanzendiversität in beweideten Grünlandgemeinschaften mit einer verringerten Unkrautabundanz sowohl bei der oberirdischen Vegetation als auch in der Bodensamenbank verbunden sein kann. Die Daten wurden in einem Weidelandexperiment gesammelt, das 1994 in Missouri, USA, etabliert wurde. Das Experiment bestand aus 15 m×15 m Probeflächen, die mit Festuca arundinacea Schreb. oder Bromus inermis Leysser als Basisarten in Mischungen von 1, 2, 3, 6 oder 8 Futterarten eingesät waren. Die Probeflächen wurden während jeder Wachstumssaison von 1998 bis 2002 stark mit Vieh beweidet. Die oberirdische Pflanzenartenzusammensetzung wurde in jeder Fläche mit einer Punktstopmethode gemessen. Bodenproben wurden 1999 und 2002 gesammelt und die Artenzusammensetzung der keimfähigen Unkrautsamen wurde in den Probeflächen bewertet, indem die Keimlinge identifiziert wurden, die in einer 8-wöchigen Periode keimten. Die Artendiversität wurde unter Verwendung verschiedener Indizes gemessen: Artenreichtum (S), Shannon–Wiener-Diversitätsindex (H^′) und Futterarten-Äquitabilität (J). Die oberirdische Unkrautartenabundanz in den Probeflächen stand in keiner Beziehung zum Futterartenreichtum (S), aber die Unkrautabundanz nahm ab, wenn die Äquitabilität (J) der ansässigen Futterarten in den Mischungen zunahm. Die Artenzusammensetzung der Mischungen könnte die Unkrautabundanz beeinflusst haben. Sowohl die Unkräuter in der Bodensamenbank, als auch in der oberirdischen Vegetation waren weniger abundant in Mischungen, die F. arundinacea enthielten, im Vergleich zu denen, die B. inermis enthielten. Obgleich Variablen wie die Futterpflanzenproduktivität möglicherweise ebenfalls die Unkrautabundanz im Weideland unterdrücken, lassen unsere Ergebnisse vermuten, dass die Aufrechterhaltung einer gleichmäßigen Mischung von Futterarten ebenfalls helfen kann, die Unkräuter zu unterdrücken.     

Correlations of phytoplankton standing crop,species diversity and dominance with physical-chemical data in a coastal salt pond

A set of environmental variables from a coastal salt pond were regressed in a stepwise fashion against the following dependent variables: phytoplankton standing crop, species diversity and dominance. Multiple variable interaction, as measured by multiple correlation coefficients yielded higher values than the coefficients obtained by regressing individual environmental variables against the dependent variables. The seasonal variation in the dependent variables could, therefore, be explained to a greater extent by the effects of multiple variables rather than single limiting factors.Financial support was supplied by a fellowship from the Jessie Smith Noyes Foundation to the senior author.     

Biomass and structure of planktonic communities along an air temperature gradient in subarctic Sweden

1. Air temperature will probably have pronounced effects on the composition of plankton communities in northern lake ecosystems, either via indirect effects on the export of essential elements from catchments or through direct effects of water temperature and the ice‐free period on the behaviour of planktonic organisms. 2. We assessed the role of temperature by comparing planktonic communities in 15 lakes along a 6 °C air temperature gradient in subarctic Sweden. 3. We found that the biomass of phytoplankton, bacterioplankton and the total planktonic biomass were positively related to air temperature, probably as a result of climatic controls on the export of nitrogen from the catchment (which affects phytoplankton biomass) and dissolved organic carbon (affecting bacterioplankton biomass). 4. The structure of the zooplankton community, and top down effects on phytoplankton, were apparently not related to temperature but mainly to trophic interactions ultimately dependent on the presence of fish in the lakes. 5. Our results suggest that air temperature regimes and long‐term warming can have strong effects on the planktonic biomass in high latitude lakes. Effects of temperature on the structure of the planktonic community might be less evident unless warming permits the invasion of fish into previous fishless lakes.     

Non-random distribution of individual genetic diversity along an environmental gradient

Improving our knowledge of the links between ecology and evolution is especially critical in the actual context of global rapid environmental changes. A critical step in that direction is to quantify how variation in ecological factors linked to habitat modifications might shape observed levels of genetic variability in wild populations. Still, little is known on the factors affecting levels and distribution of genetic diversity at the individual level, despite its vital underlying role in evolutionary processes. In this study, we assessed the effects of habitat quality on population structure and individual genetic diversity of tree swallows (Tachycineta bicolor) breeding along a gradient of agricultural intensification in southern Québec, Canada. Using a landscape genetics approach, we found that individual genetic diversity was greater in poorer quality habitats. This counter-intuitive result was partly explained by the settlement patterns of tree swallows across the landscape. Individuals of higher genetic diversity arrived earlier on their breeding grounds and settled in the first available habitats, which correspond to intensive cultures. Our results highlight the importance of investigating the effects of environmental variability on individual genetic diversity, and of integrating information on landscape structure when conducting such studies.     

云南省魔芋与玉米多样性栽培控制魔芋软腐病

在云南省富源县魔芋与玉米多样性栽培区设置田间试验,分析了魔芋软腐病的发病规律、植株的生长情况及发病植株病原菌的分离频率。结果表明:魔芋与玉米轮作是控制魔芋软腐病的主要措施,它可以显著降低魔芋软腐病的死亡率,且将发病高峰期延迟1个月,轮作条件下间种控病效果比单种提高59%,间种茎基部带菌率显著低于单种;间作体系内不同栽培行植株株高差异显著,发病率差异不显著,但行间发病植株欧文氏杆菌的分离频率存在显著差异,散叶期和换头期根部和茎基部病原菌分离频率与行间距呈正相关。因此,玉米魔芋轮作间种互作条件下对软腐病的防治效果最好,间作系统内部玉米对魔芋病原菌具有一定阻隔效应。     

Matching molecular diversity and ecophysiology of benthic cyanobacteria and diatoms in communities along a salinity gradient

The phylogenetic diversity of oxygenic phototrophic microorganisms in hypersaline microbial mats and their distribution along a salinity gradient were investigated and compared with the halotolerances of closely related cultivated strains. Segments of 16S rRNA genes from cyanobacteria and diatom plastids were retrieved from mat samples by DNA extraction and polymerase chain reaction (PCR), and subsequently analysed by denaturing gradient gel electrophoresis (DGGE). Sequence analyses of DNA from individual DGGE bands suggested that the majority of these organisms was related to cultivated strains at levels that had previously been demonstrated to correlate with characteristic salinity responses. Proportional abundances of amplified 16S rRNA gene segments from phylogenetic groupings of cyanobacteria and diatoms were estimated by image analysis of DGGE gels and were generally found to correspond to abundances of the respective morphotypes determined by microscopic analyses. The results indicated that diatoms accounted for low proportions of cells throughout, that the cyanobacterium Microcoleus chthonoplastes and close relatives dominated the communities up to a salinity of 11% and that, at a salinity of 14%, the most abundant cyanobacteria were related to highly halotolerant cultivated cyanobacteria, such as the recently established phylogenetic clusters of Euhalothece and Halospirulina . Although these organisms in cultures had previously demonstrated their ability to grow with close to optimal rates over a wide range of salinities, their occurrence in the field was restricted to the highest salinities investigated.     

Patterns of diversity in plant and soil microbial communities along a productivity gradient in a Michigan old-field

The relationship between plant diversity and productivity has received much attention in ecology, but the relationship of these factors to soil microbial communities has been little explored. The carbon resources that support soil microbial communities are primarily derived from plants, so it is likely that the soil microbial community should respond to changes in plant diversity or productivity, particularly if the plant community affects the quality or quantity of available carbon. We investigated the relationship of plant diversity and productivity to the composition of the soil microbial community along a topographic gradient in a mid-successional old-field in southwestern Michigan. Soil moisture, soil inorganic N, and plant biomass increased from the top to the base of the slope, while light at ground level decreased along this same gradient. We characterized the changes in resource levels along this gradient using an index of productivity that incorporated light levels, soil N, soil moisture, and plant biomass. Average plant species richness declined with this productivity index and there were associated compositional changes in the plant community along the gradient. The plant community shifted from predominantly low-growing perennial forbs at low productivities to perennial grasses at higher productivities. Although there was variation in the structure of the soil microbial community [as indicated by fatty acid methyl ester (FAME) profiles], changes in the composition of the soil microbial community were not correlated with plant productivity or diversity. However, microbial activity [as indicated by Biolog average well color development and substrate-induced respiration (SIR)] was positively correlated with plant productivity. The similarity between patterns of plant biomass and soil microbial activity suggests that either plant productivity is driving microbial productivity or that limiting resources for each of these two communities co-vary.     

Plant and insect diversity along a pollution gradient: understanding species richness across trophic levels

We analysed species richness of plants and true bugs (Insecta, Heteroptera) along a pollution gradient in Scots pine stands in Central Germany. As a consequence of particulate deposition, pH-values of soils increased in the vicinity of the emission source. Therefore, emission increased productivity. Species richness of plants increased with decreasing distance from emission source, and thus with increasing productivity. Similarly, species richness of herbivorous Heteroptera increased with decreasing distance from emission source, whereas, surprisingly, abundance decreased. The proportion of specialised herbivorous bug species is largest in the vicinity of the emission source. Thus, the diversity pattern of herbivores may be explained by the specialisation hypothesis and not the consumer rarity hypothesis. Species richness and abundance of carnivorous Heteroptera showed no significant trend along the gradient. Overall our data favour the bottom-up control of species diversity in the analysed system.     

Clonal diversity of Phragmites australis propagating along water depth gradient

Clonal composition of a large, expanding reed stand was studied with two genetic profiling methods. Sampling was carried out along two parallel, and two perpendicular transects at the shore of Lake Balaton, Hungary, in a 900 m long reed stand, with distances between sampling points ranging from 15 to 30 m. The four primer pairs involved in microsatellite comparisons provided a total of 45 polymorphic allelic variants that determined 61 multilocus phenotypes. Along the transects, clone number decreased towards the deep water. RAPD analysis involved eleven random decamer primers and 115 repeatably amplifying and polymorphic RAPD fragments. Applying our new data evaluation method, highly similar information was gained from RAPD investigation and the microsatellite method on the number and extension of the clones. The results demonstrated that clone competition along a water depth gradient where generative reproduction is allowed only at the lakeshore edge brings about decreasing genetic diversity irrespectively of the health status of reed stand.     

Storage and decomposition of particulate organic matter along the longitudinal gradient of an agriculturally-impacted stream

An agriculturally-impacted stream in northern Idaho was examined over a two-year period to determine seasonal and longitudinal patterns of the storage and decomposition of particulate organic matter. Biomass of benthic organic matter (BOM) was considerably less than values reported in the literature for comparable, undisturbed streams. Coarse, fine, and total benthic particulate organic matter were not correlated with parameters pertaining to stream size (e.g., stream order), but were correlated with sample site and amount of litterfall. The association of BOM with site and litterfall suggests that storage of particulate organic matter is a function of local characteristics rather than stream size. Low biomass of stored organic matter is a response to the low input of terrestrially-derived organic matter resulting from removal of climax vegetation.Leaf packs of alder, Alnus sp., were placed in the stream seasonally for 30 and 60 d. While there were significant differences for months, there was no significant difference among sites for leaf packs exposed for 30 d. Significant differences were observed among both sites and months for leaf packs exposed for 60 d; however, differences among sites accounted for only 5% of the variance. The absence of differences in decomposition of organic matter along the gradient of Lapwai Creek, despite heterogeneity of the drainage basin and availability of organic matter, may be in response to the overall low biomass of stored benthic organic matter. This study demonstrates that agricultural activity can substantially influence instream heterotrophic processes through reduced availability of organic matter and can shape community structure and ecosystem dynamics of streams flowing through agricultural drainage basins.     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.