Combined effect of virus infection and water stress on water flow and water economy in grapevines

Water limitation is one of the major threats affecting grapevine production. Thus, improving water‐use efficiency (WUE) is crucial for a sustainable viticulture industry in Mediterranean regions. Under field conditions, water stress (WS) is often combined with viral infections as those are present in major grape‐growing areas worldwide. Grapevine leafroll‐associated virus 3 (GLRaV‐3) is one of the most important viruses affecting grapevines. Indeed, the optimization of water use in a real context of virus infection is an important topic that needs to be understood. In this work, we have focused our attention on determining the interaction of biotic and abiotic stresses on WUE and hydraulic conductance (K_h) parameters in two white grapevine cultivars (Malvasia de Banyalbufar and Giró Ros). Under well‐watered (WW) conditions, virus infection provokes a strong reduction (P < 0.001) in K_petiole in both cultivars; however, K_leaf was only reduced in Malvasia de Banyalbufar. Moreover, the presence of virus also reduced whole‐plant hydraulic conductance (K_hplant) in 2013 and 2014 for Malvasia de Banyalbufar and in 2014 for Giró Ros. Thus, the effect of virus infection on water flow might explain the imposed stomatal limitation. Under WS conditions, the virus effect on K_plant was negligible, because of the bigger effect of WS than virus infection. Whole‐plant WUE (WUE_WP) was not affected by the presence of virus neither under WW nor under WS conditions, indicating that plants may adjust their physiology to counteract the virus infection by maintaining a tight stomatal control and by sustaining a balanced carbon change.     

Mycorrhizal colonisation improves fruit yield and water use efficiency in watermelon (Citrullus lanatus Thunb.) grown under well-watered and water-stressed conditions

The effect of arbuscular mycorrhizal (AM) colonisation by Glomus clarum on fruit yield and water use efficiency (WUE) was evaluated in watermelon (Citrullus lanatus) cv. Crimson Sweet F1 under field conditions. Treatments were: (1) well-watered plants without mycorrhizae (WW-M), (2) well-watered plants with mycorrhizae (WW+M), (3) water- stressed plants without mycorrhizae (WS-M) and (4) water-stressed plants with mycorrhizae (WS+M). When soil water tension readings reached –20 and –50 kPa for well-watered (WW) and water-stressed (WS) treatments, respectively, irrigation was initiated to restore the top soil to near field capacity. Water stress reduced watermelon shoot and root dry matter, fruit yield, water use efficiency but not total soluble solids (TSS) in the fruit, compared with the non-stressed treatments. Mycorrhizal plants had significantly higher biomass and fruit yield compared to nonmycorrhizal plants, whether plants were water stressed or not. AM colonisation increased WUE in both WW and WS plants. Macro- (N, P, K, Ca and Mg) and micro- (Zn, Fe and Mn) nutrient concentrations in the leaves were significantly reduced by water stress. Mycorrhizal colonisation of WS plants restored leaf nutrient concentrations to levels in WW plants in most cases. This is the first report of the mitigation of the adverse effect of water stress on yield and quality of a fruit crop.     

The Effect of Groundnut rosette assistor virus on the Agronomic Performance of Four Groundnut (Arachis hypogaea L.) Genotypes

The effect of Groundnut rosette assistor virus (GRAV), in the absence of the other two agents (Groundnut rosette virus and its satellite RNA) of the groundnut rosette disease virus complex, was evaluated on the agronomic performance of four groundnut (=peanut) genotypes (CG‐7, ICGV‐SM‐90704, JL‐24 and ICG‐12991) with different botanical characteristics. All genotypes infected with GRAV showed mild yellowing/chlorosis of leaves and the symptoms persisted throughout their growth period. ELISA absorbance values indicated lower amounts of GRAV antigen in ICGV‐SM‐90704 than in the other genotypes. The reduction in leaf area due to GRAV infection varied between 15.5% and 21.7%, whereas the plant height was decreased between 11.3% and 13.4% among the four genotypes. GRAV infection caused 28.4%, 16.9%, 21.7% and 25.5% reduction in the dry weight of haulms in CG‐7, ICGV‐SM‐90704, JL‐24 and ICG‐12991 respectively. Plants infected with GRAV showed greater reduction in seed weight in CG‐7 (52.2%), followed by JL‐24 (46.1%), ICG‐12991 (40.7%) and ICGV‐SM‐90704 (25.7%). These results provide evidence for the first time that GRAV infection, without GRV and sat RNA, affect plant growth and contribute to yield losses in groundnut.     

Nitrogen enrichment lowers Betula pendula green and yellow leaf stoichiometry irrespective of effects of elevated carbon dioxide

Deep rooting has been identified as strategy for desiccation avoidance in natural vegetation as well as in crops like rice and sorghum. The objectives of this study were to determine root morphology and water uptake of four inbred lines of tropical maize (Zea mays L.) differing in their adaptation to drought. The specific questions were i) if drought tolerance was related to the vertical distribution of the roots, ii) whether root distribution was adaptive or constitutive, and iii) whether it affected water extraction, water status, and water use efficiency (WUE) of the plant. In the main experiment, seedlings were grown to the V5 stage in growth columns (0.80 m high) under well-watered (WW) and water-stressed (WS) conditions. The depth above which 95 % of all roots were located (D₉₅) was used to estimate rooting depth. It was generally greater for CML444 and Ac7729/TZSRW (P2) compared to SC-Malawi and Ac7643 (P1). The latter had more lateral roots, mainly in the upper part of the soil column. The increase in D₉₅ was accompanied by increases in transpiration, shoot dry weight, stomatal conductance and relative water content without adverse effects on the WUE. Differences in the morphology were confirmed in the V8 stage in large boxes: CML444 with thicker (0.14 mm) and longer (0.32 m) crown roots compared to SC-Malawi. Deep rooting, drought sensitive P2 showed markedly reduced WUE, likely due to an inefficient photosynthesis. The data suggest that a combination of high WUE and sufficient water acquisition by a deep root system can increase drought tolerance.

     

Reactive oxygen metabolism in mycorrhizal and non-mycorrhizal citrus (Poncirus trifoliata) seedlings subjected to water stress

The effect of the arbuscular mycorrhizal (AM) fungus, Glomus versiforme, on growth and reactive oxygen metabolism of trifoliate orange (Poncirus trifoliata) seedlings was studied in potted plants under well-watered (WW) and water stressed (WS) conditions. Water stress significantly decreased root colonization. Shoot dry weight, plant height and stem diameter were higher in AM than in non-AM seedlings regardless of the water status. Inoculation with G. versiforme increased root dry weight and leaf number per plant of WW seedlings. There was less malondialdehyde (MDA) concentration in leaves and roots of AM seedlings, as well as lower hydrogen peroxide (H(2)O(2)) and superoxide anion radical (O(2)(-)) concentrations in AM roots under WW and WS conditions. AM inoculation did not affect the H(2)O(2) and O(2)(-) concentrations of WW leaves. Whether WS or not, AM symbiosis notably increased the guaiacol peroxidase (G-POD) activity of leaves, glutathione reductase (GR) activity of leaves and ascorbate peroxidase (APX) activity of roots. AM infection also markedly increased the APX activity of WS leaves. Soluble proteins and glutathione (GSH) in leaves and roots and ascorbate (ASC) in leaves were higher in WW AM than in WW non-AM seedlings. AM infection also enhanced the ASC and GSH contents of leaves and roots in WS seedlings. Cross-tolerance might occur in AM plants and be enhanced by AM symbiosis. Our results suggest that the increased concentrations of antioxidant enzymes and non-enzymatic antioxidants found in AM plants may serve to protect the organism against oxidative damage, enhancing drought tolerance.     

Root growth and water uptake in winter wheat under deficit irrigation

Root growth is critical for crops to use soil water under water-limited conditions. A field study was conducted to investigate the effect of available soil water on root and shoot growth, and root water uptake in winter wheat (Triticum aestivum L.) under deficit irrigation in a semi-arid environment. Treatments consisted of rainfed, deficit irrigation at different developmental stages, and adequate irrigation. The rainfed plots had the lowest shoot dry weight because available soil water decreased rapidly from booting to late grain filling. For the deficit-irrigation treatments, crops that received irrigation at jointing and booting had higher shoot dry weight than those that received irrigation at anthesis and middle grain filling. Rapid root growth occurred in both rainfed and irrigated crops from floral initiation to anthesis, and maximum rooting depth occurred by booting. Root length density and dry weight decreased after anthesis. From floral initiation to booting, root length density and growth rate were higher in rainfed than in irrigated crops. However, root length density and growth rate were lower in rainfed than in irrigated crops from booting to anthesis. As a result, the difference in root length density between rainfed and irrigated treatments was small during grain filling. The root growth and water use below 1.4 m were limited by a caliche (45% CaCO₃) layer at about 1.4 m profile. The mean water uptake rate decreased as available soil water decreased. During grain filling, root water uptake was higher from the irrigated crops than from the rainfed. Irrigation from jointing to anthesis increased seasonal evapotranspiration, grain yield, harvest index and water-use efficiency based on yield (WUE), but did not affect water-use efficiency based on aboveground biomass. There was no significant difference in WUE among irrigation treatments except one-irrigation at middle grain filling. Due to a relatively deep root system in rainfed crops, the higher grain yield and WUE in irrigated crops compared to rainfed crops was not a result of rooting depth or root length density, but increased harvest index, and higher water uptake rate during grain filling.     

Elevated auxin and reduced cytokinin contents in rootstocks improve their performance and grafting success

Plant grafting is an important technique for horticultural and silvicultural production. However, many rootstock plants suffer from undesirable lateral bud outgrowth, low grafting success rates or poor rooting. Here, we used a root‐predominant gene promoter (SbUGT) to drive the expression of a tryptophan‐2‐monooxygenase gene (iaaM) from Agrobacterium tumefaciens to increase auxin levels in tobacco. The transgenic plants, when used as a rootstock, displayed inhibited lateral bud outgrowth, enhanced grafting success rate and improved root initiation. However, root elongation and biomass of SbUGT::iaaM transgenic plants were reduced compared to those of wild‐type plants. In contrast, when we used this same promoter to drive CKX (a cytokinin degradation gene) expression, the transgenic tobacco plants displayed enhanced root elongation and biomass. We then made crosses between the SbUGT::CKX and SbUGT::iaaM transgenic plants. We observed that overexpression of the CKX gene neutralized the negative effects of auxin overproduction on root elongation. Also, the simultaneous expression of both the iaaM and CKX genes in rootstock did not disrupt normal growth and developmental patterns in wild‐type scions. Our results demonstrate that expression of both the iaaM and CKX genes predominantly in roots of rootstock inhibits lateral bud release from rootstock, improves grafting success rates and enhances root initiation and biomass.     

Extraction of P solubilizing active substances from the cell wall of groundnut roots

Groundnut can take up more phosphorus (P) from a low P soil that hardly contains plant available iron-bound P (Fe-P) as its major P form than other crops. This is considered to be caused by the presence of substances in the root cell wall (CW) that are able to solubilize P. A method for extraction of these phosphorus solubilizing active substances (PSAS) from the root CW of groundnut is described in this paper. Acid, alkaline and water extractants were used, but only a treatment with 1 M NaOH at 80 °C for 24 h was found to be appropriate to extract the PSAS from the root CW. The characteristics of the CW and the extracted CW components were compared. The P solubilizing activity of both decreased sharply after addition of Fe³⁺, whereas Ca²⁺ and Mg²⁺ had no effect. This similarity in chemical characteristics suggested that we had successfully extracted the active substances in the CW. Phosphorus-solubilizing compounds were also extracted from the root CW of other crops like soybean, pigeon pea and maize, but these other crops contained less PSAS than groundnut. Using gel permeation and anion exchange column chromatography, the CW components were purified for HPLC analysis. The HPLC analyses indicated that two common retention times for the active substances existed for all four crops. The significance of the root CW in plant P nutrition is discussed.     

Net changes of soil C stocks in two grassland soils 26 months after simulated pasture renovation including biochar addition

The use of deep‐rooting pasture species as a management practice can increase the allocation of plant carbon (C) below ground and enhance C storage. A 2‐year lysimeter trial was set up to compare changes in C stocks of soils under either deep‐ or shallow‐rooting pastures and investigate whether biochar addition below the top 10 cm could promote root growth at depth. For this i) soil ploughing at cultivation was simulated in a silt loam soil and in a sandy soil by inverting the 0 to 10 and 10‐ to 20‐cm‐depth soil layers, and a distinctive biochar (selected for each soil to overcome soil‐specific plant growth limitations) was mixed at 10 Mg ha^?1 in the buried layer, where appropriate and ii) three pasture types with contrasting root systems were grown. In the silt loam, soil inversion resulted in a general loss of C (2.0–8.1 Mg ha^?1), particularly in the buried horizon, under shallow‐rooting pastures only. The addition of a C‐rich biochar (equivalent to 7.6 Mg C ha^?1) to this soil resulted in a net C gain (21–40% over the non‐biochar treatment, P < 0.10) in the buried layer under all pastures; this overcame the loss of C in this horizon under shallow‐rooting pastures. In the sandy soil, all pastures were able to maintain soil C stocks at 10–20 cm depth over time, with minor gains of C (1.6–5.1 Mg ha^?1) for the profile. In this soil, the exposure of a skeletal‐ and nutrient‐depleted soil layer at the surface may have fostered root growth at depth. The addition of a nutrient‐rich biochar (equivalent to 3.6 Mg C ha^?1) to this soil had no apparent effect on C stocks. More research is needed to understand the mechanisms through which soil C stocks at depth are preserved.     

Genotypic variation in 'early warning signals' from roots in drying soil: intrinsic differences in ABA synthesising capacity rather than root density determines total ABA 'message' in cowpea (Vigna unguiculata L.)

In a comparison of six cowpea cultivars, we determined the variation in abscisic acid (ABA) production as an ‘early warning signal’ produced in response to drought stress. By imposing drought only to the upper 20 cm rooting zone, we compared the rates of ABA synthesis relative to (i) total root mass and (ii) inherent variation per unit root mass. We were able to relate the intensity of the stress response to these two factors, and determine which is quantitatively more important as the primary signal indicating responsiveness to drought stress. Plants were grown in 1.2 m long columns and a soil drying treatment imposed in such a way that that upper roots were in dry soil and deep roots in soil at field capacity. Relative water contents (RWC) of stressed plants were similar and not significantly different from those of well watered controls. However, roots accumulated ABA in the dehydrated zone, where root water content ranged from 10–12 g g^?1 DW. The soil moisture contents and root ‐water contents in the dry zone were similar for each of the different varieties. However, the ABA contents were significantly different in drought‐stressed (upper) roots and ranged from 7.82 nmol g^?1 DW in cv. APC 689 to 16.02 nmol g^?1 DW in cv. APC 370, such that for varieties with similar overall root weights (e.g. APC 580 and APC 540) the different ABA contents were related to the capacity for ABA synthesis. The relationship between stomatal conductance and total root ABA was assessed, with a negative relation (r= 0.90, n= 24, P= 0.05) suggesting that the intrinsic capacity of cowpea varieties for ABA synthesis could play an important role in regulating stomatal conductance in a drying soil and provide useful selection criteria for tolerance to drought stress.     

Involvement of abscisic acid and cytokinins in the senescence and remobilization of carbon reserves in wheat subjected to water stress during grain filling

This study investigated the possibility that abscisic acid (ABA) and cytokinins may mediate the effect of water deficit that enhances plant senescence and remobilization of pre‐stored carbon reserves. Two high lodging‐resistant wheat (Triticum aestivum L.) cultivars were field grown and treated with either a normal or high amount of nitrogen at heading. Well‐watered (WW) and water‐stressed (WS) treatments were imposed from 9 d post‐anthesis until maturity. Chlorophyll (Chl) and photosynthetic rate (Pr) of the flag leaves declined faster in WS plants than in WW plants, indicating that the water deficit enhanced senescence. Water stress facilitated the reduction of non‐structural carbohydrate in the stems and promoted the re‐allocation of prefixed ¹⁴C from the stems to grains, shortened the grain filling period and increased the grain filling rate. Water stress substantially increased ABA but reduced zeatin (Z) + zeatin riboside (ZR) concentrations in the stems and leaves. ABA correlated significantly and negatively, whereas Z + ZR correlated positively, with Pr and Chl of the flag leaves. ABA but not Z + ZR, was positively and significantly correlated with remobilization of pre‐stored carbon and grain filling rate. Exogenous ABA reduced Chl in the flag leaves, enhanced the remobilization, and increased grain filling rate. Spraying with kinetin had the opposite effect. The results suggest that both ABA and cytokinins are involved in controlling plant senescence, and an enhanced carbon remobilization and accelerated grain filling rate are attributed to an elevated ABA level in wheat plants when subjected to water stress.     

Improving growth, flower yield, and water relations of snapdragon (Antirhinum majus L.) plants grown under well-watered and water-stress conditions using arbuscular mycorrhizal fungi

The influence of arbuscular mycorrhizal (AM) fungus Glomus deserticola (Trappe and John) on plant growth, nutrition, flower yield, water relations, chlorophyll (Chl) contents and water-use efficiency (WUE) of snapdragon (Antirhinum majus cv. butterfly) plants were studied in potted culture under well-watered (WW) and water-stress (WS) conditions. The imposed water stress condition significantly reduced all growth parameters, nutrient contents, flower yield, water relations, and Chl pigment content and increased the electrolyte leakage of the plants comparing to those of nonstressed plants. Regardless of the WS level, the mycorrhizal snapdragon plants had significantly higher shoot and root dry mass (DM), WUE, flower yield, nutrient (P, N, K, Mg, and Ca) and Chl contents than those nonmycorrhizal plants grown both under WW or WS conditions. Under WS conditions, the AM colonization had greatly improved the leaf water potential (??_w), leaf relative water content (RWC) and reduced the leaf electrolyte leakage (EL) of the plants. Although the WS conditions had markedly increased the proline content of the leaves, this increase was significantly higher in nonmycorrhizal than in mycorrhizal plants. This suggests that AM colonization enhances the host plant WS tolerance. Values of benefit and potential dry matter for AM-root associations were highest when plants were stressed and reduced under WW conditions. As a result, the snapdragon plants showed a high degree of dependency on AM fungi which improve plant growth, flower yield, water relations particularly under WS conditions, and these improvements were increased as WS level had increased. This study confirms that AM colonization can mitigate the deleterious effect of water stress on growth and flower yield of the snapdragon ornamental plant.     

Cultivar and planting date effects on soybean root growth

To avoid late summer drought, soybean [Gylcine max (L) Merrill] producers in many southern and border states of the USA modify their cropping systems. Options include use of unadapted cultivars and changing planting dates. Because root function is important to plant health and yield, this study was conducted to determine if planting date and soybean cultivar affect root growth and distribution. Seeds of one cultivar from each of four maturity groups (MG III, IV, V, and VI) were sown in mid-April, mid-May, and mid-June in 1992 and 1993 on a Tiptonville silt loam near Portageville, MO. Root observations were performed 30 and 60 days after emergence (DAE) using a minirhizotron system. Cultivars differed for root length density (RLD) only in the 15 to 28 cm depth in 1992 and in the 15 to 28 cm and 29 to 42 cm depths in 1993, but differences were not related to maturity classification of cultivar. Average RLD was 1.02 cm^–3 for MG III and IV cultivars and 1.21 cm cm^–3 for MG V and VI cultivars. Average RLD for the mid-June planting date was 1.65 cm cm^–3 but only 0.73 cm cm^–3 for the mid-April planting date. An increase in RLD between 30 and 60 DAE occurred at all soil depths. For both years, MG V and VI cultivars produced higher yields than the MG III cultivars. Earlier than normal planting dates inhibited early root growth, but did not reduce yield. Cultivars differed only slightly for the rooting characteristics measured in this study. These rooting characteristics may not be important criteria for cultivar selection.Abbreviations MG maturity group - VCR videocassette recorder - DAE days after emergence - RLD root length density - CRLD change in root length density Contribution from the Missouri Agric. Exp. Station Journal Series Number 12, 153Contribution from the Missouri Agric. Exp. Station Journal Series Number 12, 153     

Microtensiometer technique for in situ measurement of soil matric potential and root water extraction from a sandy soil

The suitability of microtensiometers to measure the spatial variation of soil matric potential and its diurnal change was tested in a pot experiment with pearl millet (Pennisetum americanum [L.] Leeke) in a sandy soil as the soil dried out.The temporal and spatial resolution of this technique allowed precise measurement of soil matric potential and thus estimation of soil water extraction from different compartments as well as from the whole rooting zone. The technique also allowed the measurement of rehydration of plants at night and root water uptake rate per unit soil volume or per unit root length. The precision of determination of root water uptake depended greatly on the accuracy of the estimate of hydraulic conductivity, which was derived from a bare soil and might be different for a cropped soil owing to aggregation induced by the root system. A linear relationship between root length and water uptake was found (r²=0.82), irrespective of variation in soil water content between compartments and despite the variation in root age, xylem differentiation and suberin formation expected to exist between different compartments of the rooting zone. As the experiment was carried out in a range of soil matric potentials between –4 and –30 kPa, drought stress did not occur. Further information at lower soil matric potentials are required, to address questions such as the importance of soil resistance for water uptake, or which portion of the root system has to be stressed to induce hormonal signals to the shoot. The microtensiometer technique can be applied to soil matric potentials up to –80 kPa.     

Root effects on soil water and hydraulic properties

Plants can affect soil moisture and the soil hydraulic properties both directly by root water uptake and indirectly by modifying the soil structure. Furthermore, water in plant roots is mostly neglected when studying soil hydraulic properties. In this contribution, we analyze effects of the moisture content inside roots as compared to bulk soil moisture contents and speculate on implications of non-capillary-bound root water for determination of soil moisture and calibration of soil hydraulic properties. In a field crop of maize (Zea mays) of 75 cm row spacing, we sampled the total soil volumes of 0.7 m × 0.4 m and 0.3 m deep plots at the time of tasseling. For each of the 84 soil cubes of 10 cm edge length, root mass and length as well as moisture content and soil bulk density were determined. Roots were separated in 3 size classes for which a mean root porosity of 0.82 was obtained from the relation between root dry mass density and root bulk density using pycnometers. The spatially distributed fractions of root water contents were compared with those of the water in capillary pores of the soil matrix. Water inside roots was mostly below 2–5% of total soil water content; however, locally near the plant rows it was up to 20%. The results suggest that soil moisture in roots should be separately considered. Upon drying, the relation between the soil and root water may change towards water remaining in roots. Relations depend especially on soil water retention properties, growth stages, and root distributions. Gravimetric soil water content measurement could be misleading and TDR probes providing an integrated signal are difficult to interpret. Root effects should be more intensively studied for improved field soil water balance calculations. Presented at the International Conference on Bioclimatology and Natural Hazards, Pol’ana nad Detvou, Slovakia, 17–20 September 2007.     

Root cooling strongly affects diel leaf growth dynamics,water and carbohydrate relations in Ricinus communis

In laboratory and greenhouse experiments with potted plants, shoots and roots are exposed to temperature regimes throughout a 24 h (diel) cycle that can differ strongly from the regime under which these plants have evolved. In the field, roots are often exposed to lower temperatures than shoots. When the root‐zone temperature in Ricinus communis was decreased below a threshold value, leaf growth occurred preferentially at night and was strongly inhibited during the day. Overall, leaf expansion, shoot biomass growth, root elongation and ramification decreased rapidly, carbon fluxes from shoot to root were diminished and carbohydrate contents of both root and shoot increased. Further, transpiration rate was not affected, yet hydrostatic tensions in shoot xylem increased. When root temperature was increased again, xylem tension reduced, leaf growth recovered rapidly, carbon fluxes from shoot to root increased, and carbohydrate pools were depleted. We hypothesize that the decreased uptake of water in cool roots diminishes the growth potential of the entire plant – especially diurnally, when the growing leaf loses water via transpiration. As a consequence, leaf growth and metabolite concentrations can vary enormously, depending on root‐zone temperature and its heterogeneity inside pots.     

Effects of elevated CO2 concentration and water deficit on fructan metabolism in Viguiera discolor Baker

Elevated [CO₂] is suggested to mitigate the negative effects of water stress in plants; however responses vary among species. Fructans are recognised as protective compounds against drought and other stresses, as well as having a role as reserve carbohydrates. We analysed the combined effects of elevated [CO₂] and water deficit on fructan metabolism in the Cerrado species Viguiera discolor Baker. Plants were cultivated for 18 days in open‐top chambers (OTC) under ambient (～380 ppm), and high (～760 ppm) [CO₂]. In each OTC, plants were submitted to three treatments: (i) daily watering (control), (ii) withholding water (WS) for 18 days and (iii) re‐watering (RW) on day 11. Analyses were performed at time 0 and days 5, 8, 11, 15 and 18. High [CO₂] increased photosynthesis in control plants and increased water use efficiency in WS plants. The decline in soil water content was more distinct in WS 760 (WS under 760 ppm), although the leaf and tuberous root water status was similar to WS 380 plants (WS under 380 ppm). Regarding fructan active enzymes, 1‐SST activity decreased in WS plants in both CO₂ concentrations, a result consistent with the decline in photosynthesis and, consequently, in substrate availability. Under WS and both [CO₂] treatments, 1‐FFT and 1‐FEH seemed to act in combination to generate osmotically active compounds and thus overcome water deficit. The proportion of hexoses to sucrose, 1‐kestose and nystose (SKN) was higher in WS plants. In WS 760, this increase was higher than in WS 380, and was not accompanied by decreases in SKN at the beginning of the treatment, as observed in WS 380 plants. These results suggest that the higher [CO₂] in the atmosphere contributed to maintain, for a longer period, the pool of hexoses and of low DP fructans, favouring the maintenance of the water status and plant survival under drought.     

Analysis of soil moisture variations in an irrigated orchard root zone

Soil moisture and suction head in an irrigated orchard were continuously monitored by time domain reflectometry (TDR) probes and gypsum blocks, respectively, during and between successive irrigation events. On each side of the trees in the plot, two 30-cm long probes were installed vertically 10 cm below the soil surface (denoted as shallow) and another two probes were installed vertically 40 cm below the soil surface (denoted as deep). The variation in moisture content measured by the TDR probes between successive irrigation events was qualitatively divided into four stages: the first was during water application; the second initiated when irrigation stopped and the moisture content in the layer sharply decreased, mainly due to free drainage. The succeeding moderate soil-moisture decrease, caused by the simultaneous diminishing free drainage and root uptake, was the third stage. During the fourth stage, moisture depletion from the layer was solely by root uptake. The slopes of moisture content variation with time throughout this stage enabled the monitoring of water availability to the plant. The range of moisture content variations and moisture depletion rates between subsequent irrigation events was higher in the shallow (10–40 cm) than in the deeper (40–70 cm) layer. Irrigation nonuniformity and spatial variability of soil hydraulic properties contributed to the unevenness of the moisture distribution in the soil profile. However, as soon as moisture content within a layer reached field capacity, namely the free drainage had stopped, irrigation uniformity had a negligible effect on water flux to the plant roots. The measured data indicate that soil moisture is fully available to the plant as long as the momentary moisture flux from the soil bulk to the soil–root interface can replenish the moisture being depleted to supply, under non-stressed conditions, the atmospheric water demand. This flux is dominated by the local momentary value of the soil's bulk hydraulic conductivity, K _r, and it stays constant for a certain range of K _r values, controlled by the increasing root suction. A decrease in water availability to the plant appears for longer irrigation intervals as a break in the constant soil-moisture depletion rate during stage 4. This break is better correlated to a threshold K _r value than to threshold values of moisture content or suction. Therefore, it is suggested that moisture content or suction used to measure water availability or to control irrigation first be alibrated by K _r() or K _r() curves, respectively.