In vegetation stands, plants receive red to far-red ratio (R:FR) signals of varying strength from all directions. However, plant responses to variations in R:FR reflected from below have been largely ignored despite their potential consequences for plant performance. Using a heterogeneous rose canopy, which consists of bent shoots down in the canopy and vertically growing upright shoots, we quantified upward farred reflection by bent shoots and its consequences for upright shoot architecture. With a three-dimensional plant model, we assessed consequences of responses to R: FR from below for plant photosynthesis. Bent shoots reflected substantially more far-red than red light, causing reduced R:FR in light reflected upwards. Leaf inclination angles increased in upright shoots which received low R:FR reflected from below. The increased leaf angle led to an increase in simulated plant photosynthesis only when this low R:FR was reflected off their own bent shoots and not when it reflected off neighbour bent shoots. We conclude that plant response to R:FR from below is an under-explored phenomenon which may have contrasting consequences for plant performance depending on the type of vegetation or crop system. The responses are beneficial for performance only when R:FR is reflected by lower foliage of the same plants. K E Y W O R D S heterogeneous canopy, light absorption, photosynthesis, red to far-red ratio, reflection, rose (Rosa hybrida), shade-avoidance, three-dimensional plant modelling 1 | INTRODUCTION Plants have limited options to escape competitive environments during their lifetime. To optimize competitiveness and ensure survival and reproduction, plants growing in vegetation stands need to show appropriate growth responses to neighbour presence by perceiving and interpreting environmental signals. In this regard, the low red to far-red ratio (R:FR) of light reflected by neighbouring vegetation (i.e., because plant tissues preferentially absorb red-and reflect farred light) is a well-studied signal of neighbour proximity (Ballaré &

Background and Aims Shading by an overhead canopy (i.e. canopy shading) entails simultaneous changes in both photosynthetically active radiation (PAR) and red to far-red ratio (R:FR). As plant responses to PAR (e.g. changes in leaf photosynthesis) are different from responses to R:FR (e.g. changes in plant architecture), and these responses occur at both organ and plant levels, understanding plant photosynthesis responses to canopy shading needs separate analysis of responses to reductions in PAR and R:FR at different levels. Methods In a glasshouse experiment we subjected plants of woody perennial rose (Rosa hybrida) to different light treatments, and so separately quantified the effects of reductions in PAR and R:FR on leaf photosynthetic traits and plant architectural traits. Using a functional–structural plant model, we separately quantified the effects of responses in these traits on plant photosynthesis, and evaluated the relative importance of changes of individual traits for plant photosynthesis under mild and heavy shading caused by virtual overhead canopies. Key Results Model simulations showed that the individual trait responses to canopy shading could have positive and negative effects on plant photosynthesis. Under mild canopy shading, trait responses to reduced R:FR on photosynthesis were generally negative and with a larger magnitude than effects of responses to reduced PAR. Conversely, under heavy canopy shading, the positive effects of trait responses to reduced PAR became dominant. The combined effects of low-R:FR responses and low-PAR responses on plant photosynthesis were not equal to the sum of the separate effects, indicating interactions between individual trait responses. Conclusions Our simulation results indicate that under canopy shading, the relative importance of plant responses to PAR and R:FR for plant photosynthesis changes with shade levels. This suggests that the adaptive significance of plant plasticity responses to one shading factor depends on plant responses to the other.

Organ temperature and variation therein plays a key role in plant functioning and its responses to e.g. climate change. There is a strong feedback between organ, especially leaf, temperature and the climate within the canopy (canopy climate), which in turn interacts with the climate outside the canopy (ambient climate). For greenhouses, the determinants of this interplay and how they drive differences between canopy and ambient climate are poorly understood. Yet, as many experiments on both regular greenhouse crops and field crops are done in greenhouses, this is crucial to know. Therefore, we designed an experiment to quantify the differences between ambient and canopy climate and leaf temperature. A path analysis was performed to quantify the interactions between components of the greenhouse canopy-climate system. We found that with high radiation the canopy climate can be up to 5˚C cooler than the ambient climate, while for cloudy days this was only 2˚C. Canopy relative humidity (RH) was up to 25% higher compared to ambient RH. We showed that radiation is very important for these climate differences, but that this effect could be partly counteracted by turning off supplementary light (i.e. due to its indirect effects e.g. changing light distribution). Leaf temperature was substantially different, both higher and lower, from the canopy air temperature. This difference was determined by leaf area index (LAI), temperature of the heating pipe and the use of supplementary light, which all strongly influence radiation, either shortwave or thermal radiation. The difference between leaf and ambient air temperature could be decreased by decreasing the LAI or increasing the temperature of the heating pipe.

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Background and Aims The success of using bent shoots in cut-rose (Rosa hybrida) production to improve flower shoot quality has been attributed to bent shoots capturing more light and thus providing more assimilates for flower shoot growth. We aimed at quantifying this contribution of photosynthesis by bent shoots to flower shoot growth. Methods Rose plants were grown with four upright flower shoots and with no, one or three bent shoots per plant. Plant architectural traits, leaf photosynthetic parameters and organ dry weight were measured. A functional–structural plant (FSP) model of rose was used to calculate photosynthesis of upright shoots and bent shoots separately, taking into account the heterogeneous canopy structure of these plants. Key Results Bent shoots contributed to 43–53 % of total assimilated CO2 by the plant. Plant photosynthesis increased by 73 and 117 % in plants with, respectively, one and three bent shoots compared with plants without bent shoots. Upright shoot photosynthesis was not significantly affected by the presence of bent shoots. However, upright shoot dry weight increased by 35 and 59 % in plants with, respectively, one and three bent shoots compared with plants without bent shoots. The increased upright shoot dry weight was entirely due to the contribution of extra photosynthesis by bent shoots, as this was the only source that could induce differences in upright shoot growth apart from their own photosynthesis. At least 47–51 % of the photosynthesis by bent shoots was translocated to upright shoots to support their biomass increase. Conclusions Based on model simulations, we conclude that the positive effect of shoot bending on flower shoot growth and quality in cut-rose production system can almost entirely be attributed to assimilate supply from bent shoots. FSP modelling is a useful tool to quantify the contributions of photosynthesis by different parts of heterogeneous canopies.