The productive performance of intercropping.

Crop diversification has been put forward as a way to reduce the environmental impact of agriculture without penalizing its productivity. In this context, intercropping, the planned combination of two or more crop species in one field, is a promising practice. On an average, intercropping saves land compared with the component sole crops, but it remains unclear whether intercropping produces a higher yield than the most productive single crop per unit area, i.e., whether intercropping achieves transgressive overyielding. Here, we quantified the performance of intercropping for the production of grain, calories, and protein in a global meta-analysis of several production indices. The results show that intercrops outperform sole crops when the objective is to achieve a diversity of crop products on a given land area. However, when intercropping is evaluated for its ability to produce raw products without concern for diversity, intercrops on average generate a small loss in grain or calorie yield compared with the most productive sole crop (-4%) but achieve similar or higher protein yield, especially with maize/legume combinations grown at moderate N supply. Overall, although intercropping does not achieve transgressive overyielding on average, our results show that intercropping performs well in producing a diverse set of crop products and performs almost similar to the most productive component sole crop to produce raw products, while improving crop resilience, enhancing ecosystem services, and improving nutrient use efficiency. Our study, therefore, confirms the great interest of intercropping for the development of a more sustainable agricultural production, supporting diversified diets.

Empirical estimates of the mutation rate for an alphabaculovirus.

Mutation rates are of key importance for understanding evolutionary processes and predicting their outcomes. Empirical mutation rate estimates are available for a number of RNA viruses, but few are available for DNA viruses, which tend to have larger genomes. Whilst some viruses have very high mutation rates, lower mutation rates are expected for viruses with large genomes to ensure genome integrity. Alphabaculoviruses are insect viruses with large genomes and often have high levels of polymorphism, suggesting high mutation rates despite evidence of proofreading activity by the replication machinery. Here, we report an empirical estimate of the mutation rate per base per strand copying (s/n/r) of Autographa californica multiple nucleopolyhedrovirus (AcMNPV). To avoid biases due to selection, we analyzed mutations that occurred in a stable, non-functional genomic insert after five serial passages in Spodoptera exigua larvae. Our results highlight that viral demography and the stringency of mutation calling affect mutation rate estimates, and that using a population genetic simulation model to make inferences can mitigate the impact of these processes on estimates of mutation rate. We estimated a mutation rate of µ = 1×10-7 s/n/r when applying the most stringent criteria for mutation calling, and estimates of up to µ = 5×10-7 s/n/r when relaxing these criteria. The rates at which different classes of mutations accumulate provide good evidence for neutrality of mutations occurring within the inserted region. We therefore present a robust approach for mutation rate estimation for viruses with stable genomes, and strong evidence of a much lower alphabaculovirus mutation rate than supposed based on the high levels of polymorphism observed.

Functional redundancy of weed seed predation is reduced by intensified agriculture.

Intensified agriculture, a driver of biodiversity loss, can diminish ecosystem functions and their stability. Biodiversity can increase functional redundancy and is expected to stabilize ecosystem functions. Few studies, however, have explored how agricultural intensity affects functional redundancy and its link with ecosystem function stability. Here, within a continental-wide study, we assess how functional redundancy of seed predation is affected by agricultural intensity and landscape simplification. By combining carabid abundances with molecular gut content data, functional redundancy of seed predation was quantified for 65 weed genera across 60 fields in four European countries. Across weed genera, functional redundancy was reduced with high field management intensity and simplified crop rotations. Moreover, functional redundancy increased the spatial stability of weed seed predation at the field scale. We found that ecosystem functions are vulnerable to disturbances in intensively managed agroecosystems, providing empirical evidence of the importance of biodiversity for stable ecosystem functions across space.

The potential of biofortification technologies for wheat and rice to fill the nutritional Zn intake gap in China.

BACKGROUND: Zinc (Zn) deficiency in humans is of worldwide concern. The objective of this study was to investigate the Zn intake gap in Chinese adults and identify the potential role of biofortification technologies for wheat and rice, including crop nutrient management and breeding, in filling the gap. RESULTS: We use data from the China Health and Nutrition Survey in 2011 to identify food consumption patterns and dietary Zn intake of 4512 adults to define and quantify the Zn intake gap in the population. The dietary Zn intake gap of surveyed adults ranged from -0.8 to 6.53 mg day-1 across nine provinces and differences were associated with differences in food consumption patterns. Both dietary Zn intake and Zn gap for males were higher than for females. The potential of changes in five management strategies (improved nitrogen fertilization, improved phosphorus fertilization, foliar Zn fertilization, improved water management and growing varieties reaching the grain Zn breeding target) was analyzed. Breeding and foliar Zn fertilization were shown to be the two most effective management strategies that could increase dietary intake by 1.29 to 5 mg Zn day-1 dependent on sex and province. CONCLUSION: This study shows that the Zn gap varied across regions in China, with some large enough to warrant interventions. Wheat and rice as two major Zn sources could be targeted without a direct need for dietary diversification. By promoting both biofortification breeding of wheat and rice and Zn fertilization, dietary Zn intake could be enhanced to contribute to human health improvement in China. © 2023 Society of Chemical Industry.

Impact of Xylella fastidiosa subspecies pauca in European olives.

Xylella fastidiosa is the causal agent of plant diseases that cause massive economic damage. In 2013, a strain of the bacterium was, for the first time, detected in the European territory (Italy), causing the Olive Quick Decline Syndrome. We simulate future spread of the disease based on climatic-suitability modeling and radial expansion of the invaded territory. An economic model is developed to compute impact based on discounted foregone profits and losses in investment. The model projects impact for Italy, Greece, and Spain, as these countries account for around 95% of the European olive oil production. Climatic suitability modeling indicates that, depending on the suitability threshold, 95.5 to 98.9%, 99.2 to 99.8%, and 84.6 to 99.1% of the national areas of production fall into suitable territory in Italy, Greece, and Spain, respectively. For Italy, across the considered rates of radial range expansion the potential economic impact over 50 y ranges from 1.9 billion to 5.2 billion Euros for the economic worst-case scenario, in which production ceases after orchards die off. If replanting with resistant varieties is feasible, the impact ranges from 0.6 billion to 1.6 billion Euros. Depending on whether replanting is feasible, between 0.5 billion and 1.3 billion Euros can be saved over the course of 50 y if disease spread is reduced from 5.18 to 1.1 km per year. The analysis stresses the necessity to strengthen the ongoing research on cultivar resistance traits and application of phytosanitary measures, including vector control and inoculum suppression, by removing host plants.

Farmers perceive diminishing ecosystem services, but overlook dis-services in intensively used agricultural landscapes in the North China Plain.

The UN sustainable development goals ask countries to advance sustainable production methods in agriculture. While the need for a transition to sustainable agricultural production is widely felt, there is little insight into local stakeholders' perceptions regarding agroecosystem (dis)services in areas with intensive production methods. The North China Plain is an agricultural production area with intensive production systems and simplified agricultural landscapes. We conducted a survey with 267 farmers in Quzhou county in the North China Plain in 2020 to measure the perceived level of agroecosystem (dis)services supply and the changes therein between 2015 and 2020. We analyzed which explanatory factors were associated with farmers' perceptions. Provisioning services were at a high level, while the regulating and supporting ecosystem services were considered to be in low supply, as evidenced by low scores for the presence of natural enemies and earthworms, and for natural habitats such as hedgerows and windbreaks. Most of the participants did not perceive dis-services from agriculture. Differences in perception between villages with contrasting biophysical and socio-economic conditions highlight the relevance of contextualized policy development for agricultural landscape composition and configuration to manage ecosystem (dis)services.

Biodiversity and yield trade-offs for organic farming.

Organic farming supports higher biodiversity than conventional farming, but at the cost of lower yields. We conducted a meta-analysis quantifying the trade-off between biodiversity and yield, comparing conventional and organic farming. We developed a compatibility index to assess whether biodiversity gains from organic farming exceed yield losses, and a substitution index to assess whether organic farming would increase biodiversity in an area if maintaining total production under organic farming would require cultivating more land at the expense of nature. Overall, organic farming had 23% gain in biodiversity with a similar cost of yield decline. Biodiversity gain is negatively correlated to yield loss for microbes and plants, but no correlation was found for other taxa. The biodiversity and yield trade-off varies under different contexts of organic farming. The overall compatibility index value was close to zero, with negative values for cereal crops, positive for non-cereal crops, and varies across taxa. Our results indicate that, on average, the proportion of biodiversity gain is similar to the proportion of yield loss for paired field studies. For some taxa in non-cereal crops, switching to organic farming can lead to a biodiversity gain without yield loss. We calculated the overall value of substitution index and further discussed the application of this index to evaluate when the biodiversity of less intensified farming system is advantageous.

Estimating the contribution of plant traits to light partitioning in simultaneous maize/soybean intercropping.

Spatial configuration and plant phenotypic plasticity contribute to increased light capture in relay intercropping, but there is little information on whether these factors also increase light capture in simultaneous intercropping. We developed and validated a three-dimensional functional-structural plant model to simulate light capture in maize and soybean sole crops and intercrop scenarios, using species traits observed in sole crops and intercrops. The intercrop maize phenotype had 2% greater light capture than the sole crop phenotype in a pure stand. The soybean intercrop phenotype had 5-10% lower light capture than the sole crop phenotype in a pure stand. The intercrop configuration increased the light capture of maize by 29% and reduced the light capture of soybean by 42%, compared with the light capture expected from sole crops. However, intercrop configuration only marginally affected total light capture by the intercrop system (+1%). Testing of individual soybean plant traits revealed that plasticity in leaf dimensions was the main reason for differences in light capture by soybean in simulated sole crops and intercrops. The results of this study illustrate a major shift of light capture from shorter species (soybean) to the taller component (maize) in a simultaneous strip intercrop. Plastic plant traits modulate this overall effect, but only marginally.

Multidecadal, county-level analysis of the effects of land use, Bt cotton, and weather on cotton pests in China.

Long-term changes in land use, climate, and agricultural technologies may affect pest severity and management. The influences of these major drivers can only be identified by analyzing long-term data. This study examines panel data on land use, adoption of genetically modified Bacillus thuringiensis (Bt) insect-resistant cotton, weather, pest severity, and insecticide use on three major cotton pests for 51 counties in China during 1991-2015. Bt cotton had pervasive effects on the whole pest complex in cotton and its management. Adoption resulted in major reductions in insecticide use for bollworm control. The resulting restoration of aphid biological control decreased aphid severity. However, mirid bugs, which have few effective natural enemies in cotton, increased in severity with warming May and reduced insecticide spraying against bollworm. The effects of landscape on pest severity were pest specific. The severity of cotton aphid and mirid bugs decreased with higher land use diversity, but the severity of highly polyphagous cotton bollworm was unrelated to land use diversity. Shares of forest, water body, and unused land area were negatively associated with the severity of mirid bugs, whereas cotton bollworm responded positively to the shares of water body and unused land area. Farmers sprayed insecticides at mild infestation levels and responded aggressively to severe bollworm outbreaks. Findings support the usefulness of Bt-based plant resistance as a component of integrated pest management (IPM) but highlight the potential for unexpected outcomes resulting from agro-ecosystem feedback loops as well as the importance of climate.

Understanding and optimizing species mixtures using functional-structural plant modelling.

Plant species mixtures improve productivity over monocultures by exploiting species complementarities for resource capture in time and space. Complementarity results in part from competition avoidance responses that maximize resource capture and growth of individual plants. Individual organs accommodate to local resource levels, e.g. with regard to nitrogen content and photosynthetic capacity or by size (e.g. shade avoidance). As a result, the resource acquisition in time and space is improved and performance of the community as a whole is increased. Modelling is needed to unravel the primary drivers and subsequent dynamics of complementary growth responses in mixtures. Here, we advocate using functional-structural plant (FSP) modelling to analyse the functioning of plant mixtures. In FSP modelling, crop performance is a result of the behaviour of the individual plants interacting through competitive and complementary resource acquisition. FSP models can integrate the interactions between structural and physiological plant responses to the local resource availability and strength of competition, which drive resource capture and growth of individuals in species mixtures. FSP models have the potential to accelerate mixed-species plant research, and thus support the development of knowledge that is needed to promote the use of mixtures towards sustainably increasing crop yields at acceptable input levels.

Quantifying within-plant spatial heterogeneity in carbohydrate availability in cotton using a local-pool model.

Background and Aims: Within-plant spatial heterogeneity in the production of and demand for assimilates may have major implications for the formation of fruits. Spatial heterogeneity is related to organ age, but also to position on the plant. This study quantifies the variation in local carbohydrate availability for the phytomers in the same cohort using a cotton growth model that captures carbohydrate production in phytomers and carbohydrate movement between phytomers. Methods: Based on field observations, we developed a functional-structural plant model of cotton that simulates production and storage of carbohydrates in individual phytomers and transport of surplus to other phytomers. Simulated total leaf area, total above-ground dry mass, dry mass distribution along the stem, and dry mass allocation fractions to each organ at the plant level were compared with field observations for plants grown at different densities. The distribution of local carbohydrate availability throughout the plant was characterized and a sensitivity analysis was conducted regarding the value of the carbohydrate transport coefficient. Key Results: The model reproduced cotton leaf expansion and dry mass allocation across plant densities adequately. Individual leaf area was underestimated at very high plant densities. Best correspondence with measured plant traits was obtained for a value of the transport coefficient of 0.1 d-1. The simulated translocation of carbohydrates agreed well with results from C-labelling studies. Moreover, simulation results revealed the heterogeneous pattern of local carbohydrate availability over the plant as an emergent model property. Conclusions: This modelling study shows how heterogeneity in local carbohydrate production within the plant structure in combination with limitations in transport result in heterogeneous satisfaction of demand over the plant. This model provides a tool to explore phenomena in cotton that are thought to be determined by local carbohydrate availability, such as branching pattern and fruit abortion in relation to climate and crop management.

Genotype assembly, biological activity and adaptation of spatially separated isolates of Spodoptera litura nucleopolyhedrovirus.

The cotton leafworm Spodoptera litura is a polyphagous insect. It has recently made a comeback as a primary insect pest of cotton in Pakistan due to reductions in pesticide use on the advent of genetically modified cotton, resistant to Helicoverpa armigera. Spodoptera litura nucleopolyhedrovirus (SpltNPV) infects S. litura and is recognized as a potential candidate to control this insect. Twenty-two NPV isolates were collected from S. litura from different agro-ecological zones (with collection sites up to 600 km apart) and cropping systems in Pakistan to see whether there is spatial dispersal and adaptation of the virus and/or adaptation to crops. Therefore, the genetic make-up and biological activity of these isolates was measured. Among the SpltNPV isolates tested for speed of kill in 3rd instar larvae of S. litura, TAX1, SFD1, SFD2 and GRW1 were significantly faster killing isolates than other Pakistani isolates. Restriction fragment length analysis of the DNA showed that the Pakistan SpltNPV isolates are all variants of a single SpltNPV biotype. The isolates could be grouped into three genogroups (A-C). The speed of kill of genogroup A viruses was higher than in group C according to a Cox' proportional hazards analysis. Sequence analysis showed that the Pakistan SpltNPV isolates are more closely related to each other than to the SpltNPV type species G2 (Pang et al., 2001). This suggests a single introduction of SpltNPV into Pakistan. The SpltNPV-PAK isolates are distinct from Spodoptera littoralis nucleopolyhedrovirus. There was a strong correlation between geographic spread and the genetic variation of SpltNPV, and a marginally significant correlation between the latter and the cropping system. The faster killing isolates may be good candidates for biological control of S. litura in Pakistan.

Herbivore-induced plant volatiles and tritrophic interactions across spatial scales.

Herbivore-induced plant volatiles (HIPVs) are an important cue used in herbivore location by carnivorous arthropods such as parasitoids. The effects of plant volatiles on parasitoids have been well characterised at small spatial scales, but little research has been done on their effects at larger spatial scales. The spatial matrix of volatiles ('volatile mosaic') within which parasitoids locate their hosts is dynamic and heterogeneous. It is shaped by the spatial pattern of HIPV-emitting plants, the concentration, chemical composition and breakdown of the emitted HIPV blends, and by environmental factors such as wind, turbulence and vegetation that affect transport and mixing of odour plumes. The volatile mosaic may be exploited differentially by different parasitoid species, in relation to species traits such as sensory ability to perceive volatiles and the physical ability to move towards the source. Understanding how HIPVs influence parasitoids at larger spatial scales is crucial for our understanding of tritrophic interactions and sustainable pest management in agriculture. However, there is a large gap in our knowledge on how volatiles influence the process of host location by parasitoids at the landscape scale. Future studies should bridge the gap between the chemical and behavioural ecology of tritrophic interactions and landscape ecology.

Achieving Durable Resistance Against Plant Diseases: Scenario Analyses with a National-Scale Spatially Explicit Model for a Wind-Dispersed Plant Pathogen.

Genetic resistance in crops is a cornerstone of disease management in agriculture. Such genetic resistance is often rapidly broken due to selection for virulence in the pathogen population. Here, we ask whether there are strategies that can prolong the useful life of plant resistance genes. In a modeling study, we compared four deployment strategies: gene pyramiding, sequential use, simultaneous use, and a mixed strategy. We developed a spatially explicit model for France and parameterized it for the fungal pathogen Puccinia striiformis f. sp. tritici (causing wheat yellow rust) to test management strategies in a realistic spatial setting. We found that pyramiding two new resistance genes in one variety was the most durable solution only when the virulent genotype had to emerge by mutation. Deploying single-gene-resistant varieties concurrently with the pyramided variety eroded the durability of the gene pyramid. We found that continuation of deployment of varieties with broken-down resistance prolonged the useful life of simultaneous deployment of four single-gene-resistant varieties versus sequential use. However, when virulence was already present in the pathogen population, durability was low and none of the deployment strategies had effect. These results provide guidance on effective strategies for using resistance genes in crop protection practice.

Wild pollinators enhance oilseed rape yield in small-holder farming systems in China.

BACKGROUND: Insect pollinators play an important role in crop pollination, but the relative contribution of wild pollinators and honey bees to pollination is currently under debate. There is virtually no information available on the strength of pollination services and the identity of pollination service providers from Asian smallholder farming systems, where fields are small, and variation among fields is high. We established 18 winter oilseed rape (Brassica napus L.) fields along a large geographical gradient in Jiangxi province in China. In each field, oilseed rape plants were grown in closed cages that excluded pollinators and open cages that allowed pollinator access. The pollinator community was sampled by pan traps for the entire oilseed rape blooming period. RESULTS: Oilseed rape plants from which insect pollinators were excluded had on average 38% lower seed set, 17% lower fruit set and 12% lower yield per plant, but the seeds were 17% heavier, and the caged plants had 28% more flowers and 18% higher aboveground vegetative biomass than plants with pollinator access. Oilseed rape plants thus compensate for pollination deficit by producing heavier seeds and more flowers. Regression analysis indicated that local abundance and diversity of wild pollinators were positively associated with seed set and yield/straw ratio, while honey bee abundance was not related to yield parameters. CONCLUSIONS: Wild pollinator abundance and diversity contribute to oilseed rape yield by enhancing plant resource allocation to seeds rather than to above-ground biomass. This study highlights the importance of the conservation of wild pollinators to support oilseed rape production in small-holder farming systems in China.

The contribution of phenotypic plasticity to complementary light capture in plant mixtures.

Interspecific differences in functional traits are a key factor for explaining the positive diversity-productivity relationship in plant communities. However, the role of intraspecific variation attributable to phenotypic plasticity in diversity-productivity relationships has largely been overlooked. By taking a wheat (Triticum aestivum)-maize (Zea mays) intercrop as an elementary example of mixed vegetation, we show that plasticity in plant traits is an important factor contributing to complementary light capture in species mixtures. We conceptually separated net biodiversity effect into the effect attributable to interspecific trait differences and species distribution (community structure effect), and the effect attributable to phenotypic plasticity. Using a novel plant architectural modelling approach, whole-vegetation light capture was simulated for scenarios with and without plasticity based on empirical plant trait data. Light capture was 23% higher in the intercrop with plasticity than the expected value from monocultures, of which 36% was attributable to community structure and 64% was attributable to plasticity. For wheat, plasticity in tillering was the main reason for increased light capture, whereas for intercropped maize, plasticity induced a major reduction in light capture. The results illustrate the potential of plasticity for enhancing resource acquisition in mixed stands, and indicate the importance of plasticity in the performance of species-diverse plant communities.

Intercropping enhances soil carbon and nitrogen.

Intercropping, the simultaneous cultivation of multiple crop species in a single field, increases aboveground productivity due to species complementarity. We hypothesized that intercrops may have greater belowground productivity than sole crops, and sequester more soil carbon over time due to greater input of root litter. Here, we demonstrate a divergence in soil organic carbon (C) and nitrogen (N) content over 7 years in a field experiment that compared rotational strip intercrop systems and ordinary crop rotations. Soil organic C content in the top 20 cm was 4% ± 1% greater in intercrops than in sole crops, indicating a difference in C sequestration rate between intercrop and sole crop systems of 184 ± 86 kg C ha(-1) yr(-1). Soil organic N content in the top 20 cm was 11% ± 1% greater in intercrops than in sole crops, indicating a difference in N sequestration rate between intercrop and sole crop systems of 45 ± 10 kg N ha(-1) yr(-1). Total root biomass in intercrops was on average 23% greater than the average root biomass in sole crops, providing a possible mechanism for the observed divergence in soil C sequestration between sole crop and intercrop systems. A lowering of the soil δ(15) N signature suggested that increased biological N fixation and/or reduced gaseous N losses contributed to the increases in soil N in intercrop rotations with faba bean. Increases in soil N in wheat/maize intercrop pointed to contributions from a broader suite of mechanisms for N retention, e.g., complementary N uptake strategies of the intercropped plant species. Our results indicate that soil C sequestration potential of strip intercropping is similar in magnitude to that of currently recommended management practises to conserve organic matter in soil. Intercropping can contribute to multiple agroecosystem services by increased yield, better soil quality and soil C sequestration.

Shifts in dynamic regime of an invasive lady beetle are linked to the invasion and insecticidal management of its prey.

The spread and impact of invasive species may vary over time in relation to changes in the species itself, the biological community of which it is part, or external controls on the system. We investigate whether there have been changes in dynamic regimes over the last 20 years of two invasive species in the midwestern United States, the multicolored Asian lady beetle Harmonia axyridis and the soybean aphid Aphis glycines. We show by model selection that after its 1993 invasion into the American Midwest, the year-to-year population dynamics of H. axyridis were initially governed by a logistic rule supporting gradual rise to a stable carrying capacity. After invasion of the soybean aphid in 2000, food resources at the landscape level became abundant, supporting a higher year-to-year growth rate and a higher but unstable carrying capacity, with two-year cycles in both aphid and lady beetle abundance as a consequence. During 2005-2007, farmers in the Midwest progressively increased their use of insecticides for managing A. glycines, combining prophylactic seed treatment with curative spraying based on thresholds. This human intervention dramatically reduced the soybean aphid as a major food resource for H. axyridis at landscape level and corresponded to a reverse shift towards the original logistic rule for year-to-year dynamics. Thus, we document a short episode of major predator-prey fluctuations in an important agricultural system resulting from two biological invasions that were apparently damped by widespread insecticide use. Recent advances in development of plant resistance to A. glycines in soybeans may mitigate the need for pesticidal control and achieve the same stabilization of pest and predator populations at lower cost and environmental burden.

Early competition shapes maize whole-plant development in mixed stands.

Mixed cropping is practised widely in developing countries and is gaining increasing interest for sustainable agriculture in developed countries. Plants in intercrops grow differently from plants in single crops, due to interspecific plant interactions, but adaptive plant morphological responses to competition in mixed stands have not been studied in detail. Here the maize (Zea mays) response to mixed cultivation with wheat (Triticum aestivum) is described. Evidence is provided that early responses of maize to the modified light environment in mixed stands propagate throughout maize development, resulting in different phenotypes compared with pure stands. Photosynthetically active radiation (PAR), red:far-red ratio (R:FR), leaf development, and final organ sizes of maize grown in three cultivation systems were compared: pure maize, an intercrop with a small distance (25cm) between maize and wheat plants, and an intercop with a large distance (44cm) between the maize and the wheat. Compared with maize in pure stands, maize in the mixed stands had lower leaf and collar appearance rates, increased blade and sheath lengths at low ranks and smaller sizes at high ranks, increased blade elongation duration, and decreased R:FR and PAR at the plant base during early development. Effects were strongest in the treatment with a short distance between wheat and maize strips. The data suggest a feedback between leaf initiation and leaf emergence at the plant level and coordination between blade and sheath growth at the phytomer level. A conceptual model, based on coordination rules, is proposed to explain the development of the maize plant in pure and mixed stands.

Towards modelling the flexible timing of shoot development: simulation of maize organogenesis based on coordination within and between phytomers.

BACKGROUND AND AIMS: Experimental evidence challenges the approximation, central in crop models, that developmental events follow a fixed thermal time schedule, and indicates that leaf emergence events play a role in the timing of development. The objective of this study was to build a structural development model of maize (Zea mays) based on a set of coordination rules at organ level that regulate duration of elongation, and to show how the distribution of leaf sizes emerges from this. METHODS: A model of maize development was constructed based on three coordination rules between leaf emergence events and the dynamics of organ extension. The model was parameterized with data from maize grown at a low plant population density and tested using data from maize grown at high population density. KEY RESULTS: The model gave a good account of the timing and duration of organ extension. By using initial conditions associated with high population density, the model reproduced well the increase in blade elongation duration and the delay in sheath extension in high-density populations compared with low-density populations. Predictions of the sizes of sheaths at high density were accurate, whereas predictions of the dynamics of blade length were accurate up to rank 9; moderate overestimation of blade length occurred at higher ranks. CONCLUSIONS: A set of simple rules for coordinated growth of organs is sufficient to simulate the development of maize plant structure without taking into account any regulation by assimilates. In this model, whole-plant architecture is shaped through initial conditions that feed a cascade of coordination events.