Differential responses to two heatwave intensities in a Mediterranean citrus orchard are identified by combining measurements of fluorescence and carbonyl sulfide (COS) and CO2 uptake.

The impact of extreme climate episodes such as heatwaves on plants physiological functioning and survival may depend on the event intensity, which requires quantification. We unraveled the distinct impacts of intense (HW) and intermediate (INT) heatwave days on carbon uptake, and the underlying changes in the photosynthetic system, in a Mediterranean citrus orchard using leaf active (pulse amplitude modulation; PAM) and canopy level passive (sun-induced; SIF) fluorescence measurements, together with CO2 , water vapor, and carbonyl sulfide (COS) exchange measurements. Compared to normal (N) days, gross CO2 uptake fluxes (gross primary production, GPP) were significantly reduced during HW days, but only slightly decreased during INT days. By contrast, COS uptake flux and SIFA (at 760 nm) decreased during both HW and INT days, which was reflected in leaf internal CO2 concentrations and in nonphotochemical quenching, respectively. Intense (HW) heatwave conditions also resulted in a substantial decrease in electron transport rates, measured using leaf-scale fluorescence, and an increase in the fractional energy consumption in photorespiration. Using the combined proxy approach, we demonstrate a differential ecosystem response to different heatwave intensities, which allows the trees to preserve carbon assimilation during INT days but not during HW days.

Quantification of leaf-scale light energy allocation and photoprotection processes in a Mediterranean pine forest under extensive seasonal drought.

Photoprotection strategies in a Pinus halepensis Mill. forest at the dry timberline that shows sustained photosynthetic activity during 6-7 month summer drought were characterized and quantified under field conditions. Measurements of chlorophyll fluorescence, leaf-level gas exchange and pigment concentrations were made in both control and summer-irrigated plots, providing the opportunity to separate the effects of atmospheric from soil water stress on the photoprotection responses. The proportion of light energy incident on the leaf surface ultimately being used for carbon assimilation was 18% under stress-free conditions (irrigated, winter), declining to 4% under maximal stress (control, summer). Allocation of absorbed light energy to photochemistry decreased from 25 to 15% (control) and from 50% to 30% (irrigated) between winter and summer, highlighting the important role of pigment-mediated energy dissipation processes. Photorespiration or other non-assimilatory electron flow accounted for 15-20% and ~10% of incident light energy during periods of high and low carbon fixation, respectively, representing a proportional increase in photochemical energy going to photorespiration in summer but a decrease in the absolute amount of photorespiratory CO2 loss. Resilience of the leaf photochemical apparatus was expressed in the complete recovery of photosystem II (PSII) efficiency (ΦPSII) and relaxation of the xanthophyll de-epoxidation state on the diurnal cycle throughout the year, and no seasonal decrease in pre-dawn maximal PSII efficiency (Fv/Fm). The response of CO2 assimilation and photoprotection strategies to stomatal conductance and leaf water potential appeared independent of whether stress was due to atmospheric or soil water deficits across seasons and treatments. The range of protection characteristics identified provides insights into the relatively high carbon economy under these dry conditions, conditions that are predicted for extended areas in the Mediterranean and other regions due to global climate change.

Source-sink relations of sunflower plants as affected by a parasite modifies carbon allocations and leaf traits.

Sunflower broomrape (Orobanche cumana) is a root holoparasitic plant causing major damage to sunflower (Helianthus annuus L.). Parasite infection initiates source-sink relations between the parasite (sink) and the host (source), allocating carbohydrates, water and nutrients to the parasite. The primary aim of the current study was to explore responses of sunflower to broomrape parasitism, specifically to examine alternations in leaf area, leaf mass per area (LMA), mesophyll structure and root hydraulic conductivity. Leaf changes revealed modifications similar to described previously in shade adapted plants, causing larger and thinner leaves. These traits were accompanied with significantly higher root hydraulics. These changes were caused by carbohydrate depletion due to source-sink relationships between the host and parasite. An Imazapic herbicide (ALS inhibitor) was used for controlling broomrape attachments and by to investigate the plasticity of the traits found. Broomrape infected plants which were treated with Imazapic had leaves similar to non-infected plants, including mesophyll structure and carbon assimilation rates. These results demonstrated source-sink effects of broomrape which cause a low-light-like acclimation behavior which is reversible.

Phelipanche aegyptiaca parasitism impairs salinity tolerance in young leaves of tomato.

The parasite Phelipanche aegyptiaca infests tomato, a crop plant that is commonly cultivated in semi-arid environments, where tomato may be subject to salt stress. Since the relationship between the two stresses -salinity and parasitism - has been poorly investigated in tomato, the effects of P. aegyptiaca parasitism on tomato growing under moderate salinity were examined. Tomatoes were grown with regular or saline water irrigation (3 and 45 mM Cl- , respectively) in soils infested with P. aegyptiaca. The infested plants accumulated higher levels of sodium and chloride ions in the roots, shoots and leaves (old and young) under both salinity levels vs. non-infected plants. There was a positive linear correlation between P. aegyptiaca biomass and salt accumulation in young tomato leaves, and a negative linear correlation between parasite biomass and the osmotic potential of young tomato leaves. Concentrations of the osmoprotectants proline, myoinositol and sucrose were reduced in infected tomato plants, which impaired the host's osmotic adjustment ability. The sensitivity of P. aegyptiaca to salt stress was manifested as a decrease in biomass. In conclusion, P. aegyptiaca parasitism reduced the salt tolerance of tomato plants by promoting the accumulation of salts from the rhizosphere and impairing the host's osmotic adjustment ability.

The potential of the spectral 'water balance index' (WABI) for crop irrigation scheduling.

Hyperspectral sensing can detect slight changes in plant physiology, and may offer a faster and nondestructive alternative for water status monitoring. This premise was tested in the current study using a narrow-band 'water balance index' (WABI), which is based on independent changes in leaf water content (1500 nm) and the efficiency of the nonphotochemical quenching (NPQ) photo-protective mechanism (531 nm). The hydraulic, photo-protective and spectral behaviors of five important crops - grapevine, corn, tomato, pea and sunflower - were evaluated under water deficit conditions in order to associate the differences in stress physiology with WABI suitability. Rapid alterations in both leaf water content and NPQ were observed in grapevine, pea and sunflower, and were effectively captured by WABI. Apart from water status monitoring, the index was also successful in scheduling the irrigation of a vineyard, despite phenological and environmental variability. Conversely, corn and tomato displayed a relatively strict stomatal regime and/or mild NPQ responses and were, thus, unsuitable for WABI-based monitoring. WABI shows great potential for irrigation scheduling of various crops, and has a clear advantage over spectral models that focus on either of the abovementioned physiological mechanisms.

Recognition of Orobanche cumana Below-Ground Parasitism Through Physiological and Hyper Spectral Measurements in Sunflower (Helianthus annuus L.).

Broomrape (Orobanche and Phelipanche spp.) parasitism is a severe problem in many crops worldwide, including in the Mediterranean basin. Most of the damage occurs during the sub-soil developmental stage of the parasite, by the time the parasite emerges from the ground, damage to the crop has already been done. One feasible method for sensing early, below-ground parasitism is through physiological measurements, which provide preliminary indications of slight changes in plant vitality and productivity. However, a complete physiological field survey is slow, costly and requires skilled manpower. In recent decades, visible to-shortwave infrared (VIS-SWIR) hyperspectral tools have exhibited great potential for faster, cheaper, simpler and non-destructive tracking of physiological changes. The advantage of VIS-SWIR is even greater when narrow-band signatures are analyzed with an advanced statistical technique, like a partial least squares regression (PLS-R). The technique can pinpoint the most physiologically sensitive wavebands across an entire spectrum, even in the presence of high levels of noise and collinearity. The current study evaluated a method for early detection of Orobanche cumana parasitism in sunflower that combines plant physiology, hyperspectral readings and PLS-R. Seeds of susceptible and resistant O. cumana sunflower varieties were planted in infested (15 mg kg-1 seeds) and non-infested soil. The plants were examined weekly to detect any physiological or structural changes; the examinations were accompanied by hyperspectral readings. During the early stage of the parasitism, significant differences between infected and non-infected sunflower plants were found in the reflectance of near and shortwave infrared areas. Physiological measurements revealed no differences between treatments until O. cumana inflorescences emerged. However, levels of several macro- and microelements tended to decrease during the early stage of O. cumana parasitism. Analysis of leaf cross-sections revealed differences in range and in mesophyll structure as a result of different levels of nutrients in sunflower plants, manifesting the presence of O. cumana infections. The findings of an advanced PLS-R analysis emphasized the correlation between specific reflectance changes in the SWIR range and levels of various nutrients in sunflower plants. This work demonstrates potential for the early detection of O. cumana parasitism on sunflower roots using hyperspectral tools.

Thermal Time Model for Egyptian Broomrape (Phelipanche aegyptiaca) Parasitism Dynamics in Carrot (Daucus carota L.): Field Validation.

Carrot, a highly profitable crop in Israel, is severely damaged by Phelipanche aegyptiaca parasitism. Herbicides can effectively control the parasite and prevent damage, but for optimal results, knowledge about the soil-subsurface phenological stage of the parasite is essential. Parasitism dynamics models have been successfully developed for the parasites P. aegyptiaca, Orobanche cumana, and Orobanche minor in the summer crops, tomato, sunflower, and red clover, respectively. However, these models, which are based on a linear relationship between thermal time and the parasitism dynamics, may not necessarily be directly applicable to the P. aegyptiaca-carrot system. The objective of the current study was to develop a thermal time model to predict the effect of P. aegyptiaca parasitism dynamics on carrot growth. For development and validation of the models, data was collected from a temperature-controlled growth experiment and from 13 plots naturally infested with P. aegyptiaca in commercial carrot fields. Our results revealed that P. aegyptiaca development is related to soil temperature. Moreover, unlike P. aegyptiaca parasitism in sunflower and tomato, which could be predicted both a linear model, P. aegyptiaca parasitism dynamics on carrot roots required a nonlinear model, due to the wider range of growth temperatures of both the carrot and the parasite. Hence, two different nonlinear models were developed for optimizing the prediction of P. aegyptiaca parasitism dynamics. Both models, a beta function model and combined model composed of a beta function and a sigmoid curve, were able to predict first P. aegyptiaca attachment. However, overall P. aegyptiaca dynamics was described more accurately by the combined model (RMSE = 14.58 and 10.79, respectively). The results of this study will complement previous studies on P. aegyptiaca management by herbicides to facilitate optimal carrot growth and handling in fields infested with P. aegyptiaca.