Consequences of intra-canopy and top LED lighting for uniformity of light distribution in a tomato crop.

In the past decade, the potential of positioning LED lamps in between the canopy (intra-canopy) to enhance crop growth and yield has been explored in greenhouse cultivation. Changes in spatial heterogeneity of light absorption that come with the introduction of intra-canopy lighting have not been thoroughly explored. We calibrated and validated an existing functional structural plant model (FSPM), which combines plant morphology with a ray tracing model to estimate light absorption at leaflet level. This FSPM was used to visualize the light environment in a tomato crop illuminated with intra-canopy lighting, top lighting or a combination of both. Model validation of light absorption of individual leaves showed a good fit (R2 = 0.93) between measured and modelled light absorption of the canopy. Canopy light distribution was then quantified and visualized in three voxel directions by means of average absorbed photosynthetic photon flux density (PPFD) and coefficient of variation (CV) within that voxel. Simulations showed that the variation coefficient within horizontal direction was higher for intra-canopy lighting than top lighting (CV=48% versus CV= 43%), while the combination of intra-canopy lighting and top lighting yielded the lowest CV (37%). Combined intra-canopy and top lighting (50/50%) had in all directions a more uniform light absorption than intra-canopy or top lighting alone. The variation was minimal when the ratio of PPFD from intra-canopy to top lighting was about 1, and increased when this ratio increased or decreased. Intra-canopy lighting resulted in 8% higher total light absorption than top lighting, while combining 50% intra-canopy lighting with 50% top lighting, increased light absorption by 4%. Variation in light distribution was further reduced when the intra-canopy LEDs were distributed over strings at four instead of two heights. When positioning LED lamps to illuminate a canopy both total light absorption and light distribution have to be considered.

Current status and future challenges in implementing and upscaling vertical farming systems.

Vertical farming can produce food in a climate-resilient manner, potentially emitting zero pesticides and fertilizers, and with lower land and water use than conventional agriculture. Vertical farming systems (VFS) can meet daily consumer demands for nutritious fresh products, forming a part of resilient food systems-particularly in and around densely populated areas. VFS currently produce a limited range of crops including fruits, vegetables and herbs, but successful implementation of vertical farming as part of mainstream agriculture will require improvements in profitability, energy efficiency, public policy and consumer acceptance. Here we discuss VFS as multi-layer indoor crop cultivation systems, exploring state-of-the-art vertical farming and future challenges in the fields of plant growth, product quality, automation, robotics, system control and environmental sustainability and how research and development, socio-economic and policy-related institutions must work together to ensure successful upscaling of VFS to future food systems.

Enhancement of crop photosynthesis by diffuse light: quantifying the contributing factors.

BACKGROUND AND AIMS: Plants use diffuse light more efficiently than direct light. However, experimental comparisons between diffuse and direct light have been obscured by co-occurring differences in environmental conditions (e.g. light intensity). This study aims to analyse the factors that contribute to an increase in crop photosynthesis in diffuse light and to quantify their relative contribution under different levels of diffuseness at similar light intensities. The hypothesis is that the enhancement of crop photosynthesis in diffuse light results not only from the direct effects of more uniform vertical and horizontal light distribution in the crop canopy, but also from crop physiological and morphological acclimation. METHODS: Tomato (Solanum lycopersicum) crops were grown in three greenhouse compartments that were covered by glass with different degrees of light diffuseness (0, 45 and 71 % of the direct light being converted into diffuse light) while maintaining similar light transmission. Measurements of horizontal and vertical photosynthetic photon flux density (PPFD) distribution in the crop, leaf photosynthesis light response curves and leaf area index (LAI) were used to quantify each factor's contribution to an increase in crop photosynthesis in diffuse light. In addition, leaf temperature, photoinhibition, and leaf biochemical and anatomical properties were studied. KEY RESULTS: The highest degree of light diffuseness (71 %) increased the calculated crop photosynthesis by 7·2 %. This effect was mainly attributed to a more uniform horizontal (33 % of the total effect) and vertical PPFD distribution (21 %) in the crop. In addition, plants acclimated to the high level of diffuseness by gaining a higher photosynthetic capacity of leaves in the middle of the crop and a higher LAI, which contributed 23 and 13 %, respectively, to the total increase in crop photosynthesis in diffuse light. Moreover, diffuse light resulted in lower leaf temperatures and less photoinhibition at the top of the canopy when global irradiance was high. CONCLUSIONS: Diffuse light enhanced crop photosynthesis. A more uniform horizontal PPFD distribution played the most important role in this enhancement, and a more uniform vertical PPFD distribution and higher leaf photosynthetic capacity contributed more to the enhancement of crop photosynthesis than did higher values of LAI.

Crop management impacts the efficiency of quantitative trait loci (QTL) detection and use: case study of fruit load×QTL interactions.

Mapping studies using populations with introgressed marker-defined genomic regions are continuously increasing knowledge about quantitative trait loci (QTL) that correlate with variation in important crop traits. This knowledge is useful for plant breeding, although combining desired traits in one genotype might be complicated by the mode of inheritance and co-localization of QTL with antagonistic effects, and by physiological trade-offs, and feed-back or feed-forward mechanisms. Therefore, integrating advances at the genetic level with insight into influences of environment and crop management on crop performance remains difficult. Whereas mapping studies can pinpoint correlations between QTL and phenotypic traits for specific conditions, ignoring or overlooking the importance of environment or crop management can jeopardize the relevance of such assessments. Here, we focus on fruit load (a measure determining competition among fruits on one plant) and its strong modulation of QTL effects on fruit size and composition. Following an integral approach, we show which fruit traits are affected by fruit load, to which underlying processes these traits can be linked, and which processes at lower and higher integration levels are affected by fruit load (and subsequently influence fruit traits). This opinion paper (i) argues that a mechanistic framework to interpret interactions between fruit load and QTL effects is needed, (ii) pleads for consideration of the context of agronomic management when detecting QTL, (iii) makes a case for incorporating interacting factors in the experimental set-up of QTL mapping studies, and (iv) provides recommendations to improve efficiency in QTL detection and use, with particular focus on model-based marker-assisted breeding.

Parthenocarpic potential in Capsicum annuum L. is enhanced by carpelloid structures and controlled by a single recessive gene.

BACKGROUND: Parthenocarpy is a desirable trait in Capsicum annuum production because it improves fruit quality and results in a more regular fruit set. Previously, we identified several C. annuum genotypes that already show a certain level of parthenocarpy, and the seedless fruits obtained from these genotypes often contain carpel-like structures. In the Arabidopsis bel1 mutant ovule integuments are transformed into carpels, and we therefore carefully studied ovule development in C. annuum and correlated aberrant ovule development and carpelloid transformation with parthenocarpic fruit set. RESULTS: We identified several additional C. annuum genotypes with a certain level of parthenocarpy, and confirmed a positive correlation between parthenocarpic potential and the development of carpelloid structures. Investigations into the source of these carpel-like structures showed that while the majority of the ovules in C. annuum gynoecia are unitegmic and anatropous, several abnormal ovules were observed, abundant at the top and base of the placenta, with altered integument growth. Abnormal ovule primordia arose from the placenta and most likely transformed into carpelloid structures in analogy to the Arabidopsis bel1 mutant. When pollination was present fruit weight was positively correlated with seed number, but in the absence of seeds, fruit weight proportionally increased with the carpelloid mass and number. Capsicum genotypes with high parthenocarpic potential always showed stronger carpelloid development. The parthenocarpic potential appeared to be controlled by a single recessive gene, but no variation in coding sequence was observed in a candidate gene CaARF8. CONCLUSIONS: Our results suggest that in the absence of fertilization most C. annuum genotypes, have parthenocarpic potential and carpelloid growth, which can substitute developing seeds in promoting fruit development.

Quantifying abortion rates of reproductive organs and effects of contributing factors using time-to-event analysis.

Time-to-event analysis, or survival analysis, is a method to analyse the timing of events and to quantify the effects of contributing factors. We apply this method to data on the timing of abortion of reproductive organs. This abortion often depends on source and sink strength. We hypothesise that the effect of source and sink strength on abortion rate can be quantified with a statistical model, obtained via survival analysis. Flower and fruit abortion in Capsicum annuum L., observed in temperature and planting density experiments, were analysed. Increasing the source strength as well as decreasing the sink strength decreased the abortion rate. The effect was non-linear, e.g. source strengths above 6g CH2O per plant per d did not decrease abortion rates further. The maximum abortion rate occurred around 100 degree-days after anthesis. Analyses in which sink strength was replaced with the number of fruits in a specified age category had an equal or better fit to the data. We discuss the advantages and disadvantages of using survival analyses for this kind of data. The technique can also be used for other crops showing reproductive organ abortion (e.g. soybean (Glycine max L.), cucumber (Cucumis sativus L.)), but also on other event types like bud break or germination.

Simulation of fruit-set and trophic competition and optimization of yield advantages in six Capsicum cultivars using functional-structural plant modelling.

BACKGROUND AND AIMS: Many indeterminate plants can have wide fluctuations in the pattern of fruit-set and harvest. Fruit-set in these types of plants depends largely on the balance between source (assimilate supply) and sink strength (assimilate demand) within the plant. This study aims to evaluate the ability of functional-structural plant models to simulate different fruit-set patterns among Capsicum cultivars through source-sink relationships. METHODS: A greenhouse experiment of six Capsicum cultivars characterized with different fruit weight and fruit-set was conducted. Fruit-set patterns and potential fruit sink strength were determined through measurement. Source and sink strength of other organs were determined via the GREENLAB model, with a description of plant organ weight and dimensions according to plant topological structure established from the measured data as inputs. Parameter optimization was determined using a generalized least squares method for the entire growth cycle. KEY RESULTS AND CONCLUSIONS: Fruit sink strength differed among cultivars. Vegetative sink strength was generally lower for large-fruited cultivars than for small-fruited ones. The larger the size of the fruit, the larger variation there was in fruit-set and fruit yield. Large-fruited cultivars need a higher source-sink ratio for fruit-set, which means higher demand for assimilates. Temporal heterogeneity of fruit-set affected both number and yield of fruit. The simulation study showed that reducing heterogeneity of fruit-set was obtained by different approaches: for example, increasing source strength; decreasing vegetative sink strength, source-sink ratio for fruit-set and flower appearance rate; and harvesting individual fruits earlier before full ripeness. Simulation results showed that, when we increased source strength or decreased vegetative sink strength, fruit-set and fruit weight increased. However, no significant differences were found between large-fruited and small-fruited groups of cultivars regarding the effects of source and vegetative sink strength on fruit-set and fruit weight. When the source-sink ratio at fruit-set decreased, the number of fruit retained on the plant increased competition for assimilates with vegetative organs. Therefore, total plant and vegetative dry weights decreased, especially for large-fruited cultivars. Optimization study showed that temporal heterogeneity of fruit-set and ripening was predicted to be reduced when fruits were harvested earlier. Furthermore, there was a 20 % increase in the number of extra fruit set.

Genetic differences in fruit-set patterns are determined by differences in fruit sink strength and a source : sink threshold for fruit set.

BACKGROUND AND AIMS: Fruit set in indeterminate plant species largely depends on the balance between source and sink strength. Plants of these species show fluctuations in fruit set during the growing season. It was tested whether differences in fruit sink strength among the cultivars explained the differences in fruit-set patterns. METHODS: Capsicum was chosen as a model plant. Six cultivars with differences in fruit set, fruit size and plant growth were evaluated in a greenhouse experiment. Fruit-set patterns, generative and vegetative sink strength, source strength and the source : sink ratio at fruit set were determined. Sink strength was quantified as potential growth rate. Fruit set was related to total fruit sink strength and the source : sink ratio. The effect of differences observed in above-mentioned parameters on fruit-set patterns was examined using a simple simulation model. KEY RESULTS: Sink strengths of individual fruits differed greatly among cultivars. Week-to-week fruit set in large-fruited cultivars fluctuated due to large fluctuations in total fruit sink strength, but in small-fruited cultivars, total fruit sink strength and fruit set were relatively constant. Large variations in week-to-week fruit set were correlated with a low fruit-set percentage. The source : sink threshold for fruit set was higher in large-fruited cultivars. Simulations showed that within the range of parameter values found in the experiment, fruit sink strength and source : sink threshold for fruit set had the largest impact on fruit set: an increase in these parameters decreased the average percentage fruit set and increased variation in weekly fruit set. Both were needed to explain the fruit-set patterns observed. The differences observed in the other parameters (e.g. source strength) had a lower effect on fruit set. CONCLUSIONS: Both individual fruit sink strength and the source : sink threshold for fruit set were needed to explain the differences observed between fruit-set patterns of the six cultivars.

Flower and fruit abortion in sweet pepper in relation to source and sink strength.

Source strength (assimilate supply) and sink strength (assimilate demand) of the plant were varied in different ways to investigate to what extent flower/fruit abortion in sweet pepper (Capsicum annuum L.) is determined by the availability of assimilates. Source strength was varied by changing the light level, plant density, and leaf pruning. Sink strength was varied by changing the temperature and the number and position of earlier formed fruits. Shading as well as heating for short periods showed that flowers/fruits were the most susceptible to abortion during the first week after anthesis. The different experiments where source strength was varied all showed that when source strength decreased, the rate of abortion increased linearly, whether source strength was decreased by shading, high plant density, or leaf pruning. That flower and fruit abortion not only depends on the source strength but also on the sink strength of competing organs is shown by varying the number or the position of earlier formed fruits. With the same source strength, the rate of abortion showed a close relationship with the growth rate of the earlier formed competing fruits, suggesting that the induction of abortion by earlier formed fruits is due to their sink strength. Most of the variation in abortion could be related to differences in vegetative growth rate, the latter being an indicator of the source-sink ratio. However, with the same vegetative growth rate, the rate of abortion was lower for the leaf pruning treatments where no competing fruits were retained than for the fruit load treatments. This indicates that although most of the variation in abortion can be related to the source and sink strength of the plant, some effects of competing fruits can only be explained by a combination of competition and dominance.

Simulation of leaf area development based on dry matter partitioning and specific leaf area for cut chrysanthemum.

This work aims to predict time courses of leaf area index (LAI) based on dry matter partitioning into the leaves and on specific leaf area of newly formed leaf biomass (SLA(n)) for year-round cut chrysanthemum crops. In five glasshouse experiments, each consisting of several plant densities and planted throughout the year, periodic destructive measurements were conducted to develop empirical models for partitioning and for SLA(n). Dry matter partitioning into leaves, calculated as incremental leaf dry mass divided by incremental shoot dry mass between two destructive harvests, could be described accurately (R(2 )= 0.93) by a Gompertz function of relative time, R(t). R(t) is 0 at planting date, 1 at the start of short-days, and 2 at final harvest. SLA(n), calculated as the slope of a linear regression between periodic measurements of leaf dry mass (LDM) and LAI, showed a significant linear increase with the inverse of the daily incident photosynthetically active radiation (incident PAR, MJ m(-2 )d(-1)), averaged over the whole growing period, the average glasshouse temperature and plant density (R(2 )= 0.74). The models were validated by two independent experiments and with data from three commercial growers, each with four planting dates. Measured shoot dry mass increase, initial LAI and LDM, plant density, daily temperature and incident PAR were input into the model. Dynamics of LDM and LAI were predicted accurately by the model, although in the last part of the cultivation LAI was often overestimated. The slope of the linear regression of simulated against measured LDM varied between 0.95 and 1.09. For LAI this slope varied between 1.01 and 1.12. The models presented in this study are important for the development of a photosynthesis-driven crop growth model for cut chrysanthemum crops.

Modelling of temperature-controlled internode elongation applied to chrysanthemum.

The DIF concept states that equal internode length can be achieved with the same difference between day and night temperature irrespective of the mean 24 h temperature. However, the physiological background of the DIF concept is unclear. An attempt to model internode elongation is presented based on three plausible processes, namely (1) the accumulation of elongation requirements during the day, (2) elongation during the night using elongation requirements and (3) the limitation of internode length due to low turgor pressure unable to counter cell wall elasticity. Each reaction rate constant, one per process, depends on temperature according to Arrhenius' Law. The resulting process-based model describes internode elongation in time and was calibrated on a chrysanthemum data set. Chrysanthemum plants were grown in growth chambers with rigorously defined day and night temperatures. In total, 16 temperature treatments were applied, resulting from the combination of four day and four night temperatures (16, 20, 24 and 28 degrees C). Internode elongation was measured for the tenth internode in ten plants per treatment. The percentage variance accounted for, R2adj, was almost 91%. Transferability of model parameters was shown to exist by cross validation. Simulation of the internode length in time as function of mean 24 h temperature and DIF showed that the DIF concept is not apparent after a growing period of 10 d, but is visible after 20 d. This model structure for describing internode elongation might also be applicable for other plants that show the DIF concept.

Effect of day and night temperature on internode and stem length in chrysanthemum: is everything explained by DIF?

In many plant species, including chrysanthemum, a strong positive correlation between internode length and DIF [difference between day (DT) and night (NT) temperature] has been observed. However, Langton and Cockshull (1997. Scientia Horticulturae 69: 229-237) reported no such relationship and showed that absolute DT and NT explained internode length rather than DIF. To investigate these conflicting results and to clarify the validity of the DIF concept, cut chrysanthemums (Chrysanthemum 'Reagan Improved') were grown in growth chambers at all 16 combinations of four DT and four NT (16, 20, 24 and 28 degrees C) with a 12 h day length. Length of internode 10, number of internodes and stem length were measured on days 5, 10, 17, 22 and 27 after starting the temperature treatments. Internode length on day 10 showed a positive linear relationship with DIF (R2 = 0.64). However, when internodes had reached their final length in all treatments (day 27), a much stronger positive linear relation was observed (R2 = 0.81). A model to predict final internode length was developed based on the absolute DT and NT responses: both responses were optimum curves and no significant interaction between DT and NT occurred [final internode length (mm) = -32.23 + 3.56DT + 1.08NT - 0.0687DT2 - 0.0371NT2; R2 = 0.91, where TD is day temperature and TN is night temperature]. It is shown that DIF can predict final internode length only within a temperature range where effects of DT and NT are equal in magnitude and opposite in sign (18-24 degrees C). Internode appearance rate, as well as stem length formed during the experiment, showed an optimum response to DT.