Repeated heat stress events during the reproductive phase impact the dynamic development of seeds in Brassica napus L.

Many studies pointed out the deleterious effects of high temperatures events during the crop reproductive phase on seed yield and quality. However, plant responses to repeated stressing events remain poorly understood, while the increased frequency of extreme abiotic constraints, such as spring and summer heat waves, has been proven as one feature of the on-going and future climate change. The responses of oilseed rape plants subjected to three heat stress sequences that differed in the intensity, the timing of application, the duration and the frequency of the high temperature events were investigated throughout the seed development and maturation phases under controlled conditions. Seed yield and components were measured in three different harvest dates. Biochemical and histological analyses of seeds were carried out in order to monitor the evolution of the main storage compounds (fatty acids, proteins, sugars) involved in seed nutritional quality. Although the effects of heat stress were not significant on total yield, differences in seed number and weight highlighted the strong compensation capacity in indeterminate growth species. Heat stress induced significant decreases and increases in seed oil and protein content respectively, to different extent according to the age of the pods. Soluble sugars concentrations were impacted by heat during seed development, but not when the seeds reached physiological maturity, thus indicating compensatory mechanisms that set up after the stress exposure. Our results led to conclude that the effects of repeated heat stresses on seed yield and quality were tightly related to (i) the optimal temperature of a given compound biosynthesis process, and (ii) the synchrony between the temperature event and the period of biosynthesis of the targeted storage compound. These results highlight the complexity to design thermo-sensitizing protocols to maintain or even improve the various seed quality related criteria, especially in species with indeterminate growth.

The Dynamics of Phosphorus Uptake and Remobilization during the Grain Development Period in Durum Wheat Plants.

Post-anthesis phosphorus (P) uptake and the remobilization of the previously acquired P are the principal sources of grain P nutrition in wheat. However, how the acquired P reaches the grains and its partitioning at the whole plant level remain poorly understood. Here, the temporal dynamics of the newly acquired P in durum wheat organs and its allocation to grain were examined using pulse-chase 32P-labeling experiments at 5 and 14 days after anthesis. Durum wheat plants were grown hydroponically under high and low P supplies. Each labeling experiment lasted for 24 h. Plants were harvested 24, 48, and 96 h after labeling. Low and high P treatments significantly affected the allocation of the newly acquired P at the whole plant level. Three days (96 h) after the first 32P-labeling, 8% and 4% of the newly acquired P from exogenous solution were allocated to grains, 73% and 55% to the remainder aboveground organs, and 19% and 41% to the roots at low and high P supplies, respectively. Three days after the second labeling, the corresponding values were 48% and 20% in grains, 44% and 53% in the remainder aboveground organs, and 8% and 27% in roots at low and high P supplies, respectively. These results reveal that the dynamics of P allocation to grain was faster in plants grown under low P supply than under high supply. However, the obtained results also indicate that the origin of P accumulated in durum wheat grains was mainly from P remobilization with little contribution from post-anthesis P uptake. The present study emphasizes the role of vegetative organs as temporary storage of P taken up during the grain filling period before its final allocation to grains.

Understanding the Adaptive Mechanisms of Plants to Enhance Phosphorus Use Efficiency on Podzolic Soils in Boreal Agroecosystems.

Being a macronutrient, phosphorus (P) is the backbone to complete the growth cycle of plants. However, because of low mobility and high fixation, P becomes the least available nutrient in podzolic soils; hence, enhancing phosphorus use efficiency (PUE) can play an important role in different cropping systems/crop production practices to meet ever-increasing demands in food, fiber, and fuel. Additionally, the rapidly decreasing mineral phosphate rocks/stocks forced to explore alternative resources and methods to enhance PUE either through improved seed P reserves and their remobilization, P acquisition efficiency (PAE), or plant's internal P utilization efficiency (IPUE) or both for sustainable P management strategies. The objective of this review article is to explore and document important domains to enhance PUE in crop plants grown on Podzol in a boreal agroecosystem. We have discussed P availabilities in podzolic soils, root architecture and morphology, root exudates, phosphate transporters and their role in P uptake, different contributors to enhance PAE and IPUE, and strategies to improve plant PUE in crops grown on podzolic soils deficient in P and acidic in nature.

Genetic Variation in Root Architectural Traits in Lactuca and Their Roles in Increasing Phosphorus-Use-Efficiency in Response to Low Phosphorus Availability.

Low phosphorus (P) bioavailability in the soil and concerns over global P reserves have emphasized the need to cultivate plants that acquire and use P efficiently. Root architecture adaptation to low P can be variable depending on species or even genotypes. To assess the genetic variability of root architectural traits and their responses to low P in the Lactuca genus, we examined fourteen genotypes including wild species, ancient and commercial lettuce cultivars at low (LP, 0.1 mmol. L-1) and high P (HP, 1 mmol. L-1). Plants were grown in cylindrical pots adapted for the excavation and observation of root systems, with an inert substrate. We identified substantial genetic variation in all the investigated root traits, as well as an effect of P availability on these traits, except on the diameter of thinner roots. At low P, the main responses were a decrease in taproot diameter, an increase in taproot dominance over its laterals and an increase in the inter-branch distance. Although the genotype x P treatment effect was limited to root depth, we identified a tradeoff between the capacity to maintain a thick taproot at low P and the dominance of the taproot over its laterals. Regardless of the P level, the phosphorus-use-efficiency (PUE) varied among lettuce genotypes and was significantly correlated with total root biomass regardless of the P level. As taproot depth and maximum apical diameter were the principal determinants of total root biomass, the relative increase in PUE at low P was observed in genotypes that showed the thickest apical diameters and/or those whose maximal apical diameter was not severely decreased at low P availability. This pre-eminence of the taproot in the adaptation of Lactuca genotypes to low P contrasts with other species which rely more on lateral roots to adapt to P stress.

Scion genotypes exert long distance control over rootstock transcriptome responses to low phosphate in grafted grapevine.

BACKGROUND: Grafting is widely used in horticulture and rootstocks are known to modify scion growth and adaptation to soil conditions. However, the role of scion genotype in regulating rootstock development and functioning has remained largely unexplored. In this study, reciprocal grafts of two grapevine genotypes were produced as well as the corresponding homo-graft controls. These plants were subjected to a low phosphate (LP) treatment and transcriptome profiling by RNA sequencing was done on root samples collected 27 h after the onset of the LP treatment. RESULTS: A set of transcripts responsive to the LP treatment in all scion/rootstock combinations was identified. Gene expression patterns associated with genetic variation in response to LP were identified by comparing the response of the two homo-grafts. In addition, the scion was shown to modify root transcriptome responses to LP in a rootstock dependent manner. A weighted gene co-expression network analysis identified modules of correlated genes; the analysis of the association of these modules with the phosphate treatment, and the scion and rootstock genotype identified potential hub genes. CONCLUSIONS: This study provides insights into the response of grafted grapevine to phosphate supply and identifies potential shoot-to-root signals that could vary between different grapevine genotypes.

Contribution of External and Internal Phosphorus Sources to Grain P Loading in Durum Wheat (Triticum durum L.) Grown Under Contrasting P Levels.

Phosphorus (P) in durum wheat grains after anthesis originates from either the external P source or the internal remobilization of P from different plant organs. The supply of P and its use by the plant are important factors that can affect the contribution of each source to grain P nutrition. Thus, this experiment aimed to quantify the origin of P in grains of durum wheat plants with different P nutritional status. Wheat plants were grown from juvenile stages to maturity in complete nutrient solutions with either high (0.125 mM) or low (0.025 mM) P concentrations in greenhouse conditions. Phosphorus in nutrient solutions was spiked by introducing 32P after anthesis to quantify the external P uptake and its partitioning within plant organs (spikelets, leaves, stems, roots, and post-anthesis tillers) and grains. Phosphorus use efficiency in durum wheat plants was also determined. The low and high P supply resulted in two highly different plant nutritional P status. Plants with low P status remobilized most of their stored P in all organs and allocated more than 72% of post-anthesis P uptake to grain P nutrition, whereas in the high P plants this was only 56%. Enhanced remobilization of P and the efficient allocation of newly acquired P to grains were crucial for durum wheat grain P nutrition grown under low P supply. The remobilization of P represented 81% of grain P in low P plants while it represented 65% for high P plants. Organs that contributed the most to P remobilization in low P plants were spikelets (43%) and leaves (35%). The post-anthesis tiller development was reduced in low P plants suggesting a preferential allocation of P to grains under this treatment. We concluded that P loading into grains in durum wheat is mainly derived from the remobilization of internal P sources stored before anthesis, even at high external P supply during grain filling.

Carbon and Phosphorus Allocation in Annual Plants: An Optimal Functioning Approach.

Phosphorus (P) is the second most important nutrient after nitrogen (N) and can greatly diminish plant productivity if P supply is not adequate. Plants respond to soil P availability by adjusting root biomass to maintain uptake and productivity due to P use. In spite of our vast knowledge on P effects on plant growth, how to functionally model enhanced root biomass allocation in low P environments is not fully explored. We develop a dynamic plant model based on the principle of optimal carbon (C) and P allocation to investigate growth and functional response to contrasting levels of soil P availability. By describing plant growth as a balance of growth and respiration processes, we optimize C and P allocation in order to maximize leaf productivity and drive plant response. We compare our model to a field trial and a set of hydroponic experiments which describe plant response at varying P availabilities. The model is able to reproduce long-term plant functional response to different P levels like change in root-shoot ratio (RSR), total biomass and organ P concentration. But it is not capable of fully describing the time evolution of organ P uptake and cycling within the plant. Most notable is the underestimation of organ P uptake during the vegetative growth stage which is due to the model's leaf productivity formalism. In spite of the model's parsimonious nature, which optimizes for and predicts whole plant response through leaf productivity alone, the optimal growth hypothesis can provide a reasonable framework for modelling plant response to environmental change that can be used in more physically driven vegetation models.

Importance of the vegetation-groundwater-stream continuum to understand transformation of biogenic carbon in aquatic systems - A case study based on a pine-maize comparison in a lowland sandy watershed (Landes de Gascogne, SW France).

During land-aquatic transfer, carbon (C) and inorganic nutrients (IN) are transformed in soils, groundwater, and at the groundwater-surface water interface as well as in stream channels and stream sediments. However, processes and factors controlling these transfers and transformations are not well constrained, particularly with respect to land use effect. We compared C and IN concentrations in shallow groundwater and first-order streams of a sandy lowland catchment dominated by two types of land use: pine forest and maize cropland. Contrary to forest groundwater, crop groundwater exhibited oxic conditions all-year round as a result of higher evapotranspiration and better lateral drainage that decreased the water table below the organic-rich soil horizon, prevented the leaching of soil-generated dissolved organic carbon (DOC) in groundwater, and thus limited consumption of dissolved oxygen (O2). In crop groundwater, oxic conditions inhibited denitrification and methanogenesis resulting in high nitrate (NO3-; on average 1140 ±â€¯485 µmol L-1) and low methane (CH4; 40 ±â€¯25 nmol L-1) concentrations. Conversely, anoxic conditions in forest groundwater led to lower NO3- (25 ±â€¯40 µmol L-1) and higher CH4 (1770 ±â€¯1830 nmol L-1) concentrations. The partial pressure of carbon dioxide (pCO2; 30,650 ±â€¯11,590 ppmv) in crop groundwater was significantly lower than in forest groundwater (50,630 ±â€¯26,070 ppmv), and was apparently caused by the deeper water table delaying downward diffusion of soil CO2 to the water table. In contrast, pCO2 was not significantly different in crop (4480 ±â€¯2680 ppmv) and forest (4900 ±â€¯4500 ppmv) streams, suggesting faster degassing in forest streams resulting from greater water turbulence. Although NO3-concentrations indicated that denitrification occurred in riparian-forest groundwater, crop streams nevertheless exhibited important signs of spring and summer eutrophication such as the development of macrophytes. Stream eutrophication favored development of anaerobic conditions in crop stream sediments, as evidenced by increased ammonia (NH4+) and CH4 in stream waters and concomitant decreased in NO3- concentrations as a result of sediment denitrification. In crop streams, dredging and erosion of streambed sediments during winter sustained high concentration of particulate organic C, NH4+ and CH4. In forest streams, dissolved iron (Fe2+), NH4+ and CH4 were negatively correlated with O2 reflecting the gradual oxygenation of stream water and associated oxidations of Fe2+, NH4+ and CH4. The results overall showed that forest groundwater behaved as source of CO2 and CH4 to streams, the intensity depending on the hydrological connectivity among soils, groundwater, and streams. CH4 production was prevented in cropland in soils and groundwater, however crop groundwater acted as a source of CO2 to streams (but less so than forest groundwater). Conversely, in streams, pCO2 was not significantly affected by land use while CH4 production was enhanced by cropland. At the catchment scale, this study found substantial biogeochemical heterogeneity in C and IN concentrations between forest and crop waters, demonstrating the importance of including the full vegetation-groundwater-stream continuum when estimating land-water fluxes of C (and nitrogen) and attempting to understand their spatial and temporal dynamics.

Sensitivity analyses for improving sulfur management strategies in winter oilseed rape.

Because sulfur (S) depletion in soil results in seed yield losses and grain quality degradation, especially in high S-demanding crops such as oilseed rape (Brassica napus L.), monitoring S fertilisation has become a central issue. Crop models can be efficient tools to conduct virtual experiments under different fertilisation management strategies. Using the process-based model SuMoToRI, we aimed to analyse the impact of different S fertilisation strategies coupled with the variablility observed in major plant characteristics in oilseed rape i.e. radiation use efficiency (RUE), carbon (C) allocation to the leaves (ß) and specific leaf area (SLA) on plant performance-driven variables encompassing total biomass (TDW), S in the photosynthetic leaves (QSmobile.GL) and leaf area index (LAIGL). The contrasting S supply conditions differed in the amount of S (5 levels), and the timing of application (at bolting and/or at flowering, which included a fractioned condition). For this purpose, we performed a global sensitivity analysis (GSA) and calculated two sensitivity indices i.e. the Partial Raw Correlation Coefficient (PRCC) and the Sobol index. The results showed that whatever the timing of S supply, TDW, LAIGL and QSmobile.GL increased as S input increased. For a given S supply, there was no difference in TDW, LAIGL and QSmobile.GL between a single and a fractioned supply. Moreover, delaying the supply until flowering reduced the TDW and LAIGL whereas QSmobile.GL increased. Results showed that RUE had the greatest impact on TDW under all levels of S supply and all application timings, followed by ß and SLA. RUE mostly impacted on QSmobile.GL, depending on S supply conditions, whereas it was the parameter with the least impact on LAIGL. Ultimately, our results provide strong evidence of optimised S fertilisation timings and plant characteristics that will guide producers in their agricultural practices by using specific varieties under constrained S fertilisation strategies.

Phosphorus acquisition efficiency and phosphorus remobilization mediate genotype-specific differences in shoot phosphorus content in grapevine.

Crop productivity is limited by phosphorus (P) and this will probably increase in the future. Rootstocks offer a means to increase the sustainability and nutrient efficiency of agriculture. It is known that rootstocks alter petiole P concentrations in grapevine. The objective of this work was to determine which functional processes are involved in genotype-specific differences in scion P content by quantifying P uptake, P remobilization from the reserves in the cutting and P allocation within the plant in three grapevine genotypes. Cuttings of two American rootstocks and one European scion variety were grown in sand and irrigated with a nutrient solution containing either high P (0.6 mM) or low P (0 mM). The high P solution was labelled with 32P throughout the experiment. The grapevine genotypes studied show variation in the inhibition of shoot and root biomass in response to low P supply, and P supply also affected shoot, but not root, P concentrations. Genotype-specific differences in total P content were related to differences in P acquisition and utilization efficiencies (PAE and PUE, respectively). Phosphorus allocation within the plant was not affected by genotype or P supply. The rootstock genotype known to confer high petiole P content in the vineyard was associated with a high PAE under high P, and a high PUE under low P. This suggests that the petiole P concentrations in the vineyard are related to genotype-specific differences in PAE and PUE, and that these traits could be used for rootstock selection programmes in the future.

SuMoToRI, an Ecophysiological Model to Predict Growth and Sulfur Allocation and Partitioning in Oilseed Rape (Brassica napus L.) Until the Onset of Pod Formation.

Sulfur (S) nutrition in rapeseed (Brassica napus L.) is a major concern for this high S-demanding crop, especially in the context of soil S oligotrophy. Therefore, predicting plant growth, S plant allocation (between the plant's compartments) and S pool partitioning (repartition of the mobile-S vs. non-mobile-S fractions) until the onset of reproductive phase could help in the diagnosis of S deficiencies during the early stages. For this purpose, a process-based model, SuMoToRI (Sulfur Model Toward Rapeseed Improvement), was developed up to the onset of pod formation. The key features rely on (i) the determination of the S requirements used for growth (structural and metabolic functions) through critical S dilution curves and (ii) the estimation of a mobile pool of S that is regenerated by daily S uptake and remobilization from senescing leaves. This study describes the functioning of the model and presents the model's calibration and evaluation. SuMoToRI was calibrated and evaluated with independent datasets from greenhouse experiments under contrasting S supply conditions. It is run with a small number of parameters with generic values, except in the case of the radiation use efficiency, which was shown to be modulated by S supply. The model gave satisfying predictions of the dynamics of growth, S allocation between compartments and S partitioning, such as the mobile-S fraction in the leaves, which is an indicator of the remobilization potential toward growing sinks. The mechanistic features of SuMoToRI provide a process-based framework that has enabled the description of the S remobilizing process in a species characterized by senescence during the vegetative phase. We believe that this model structure could be useful for modeling S dynamics in other arable crops that have similar senescence-related characteristics.