Comparison of continuously recorded plant-based water stress indicators for young lemon trees

Continuously recorded plant-based water stress indicators (sap flow and parameters derived from trunk diameter fluctuations) were compared in potted young lemon trees (Citrus limon (L.) Burm. fil, cv. Verna) grafted on sour orange (C. aurantium L.) rootstock submitted to deficit irrigation. Daily maximum (MXTD) and minimum (MNTD) trunk diameters and daily trunk diameter shrinkage (MDS) were directly influenced by the water supply to the trees from the soil. The continuously recorded plant-based water stress indicators presented different degrees of sensitivity when used to estimate the water status of the plants. Sap flow (SF) and MDS were more immediate and sensitive than MXTD and MNTD. However, the higher signal intensity: noise for SF and the fact that its signal intensities remained clearly above unity during the stress period, indicating that the soil water was depleted, point to the greater reliability of this indicator. Also, the possibility of developing further baseline relationships between SF and air vapour pressure deficit in fully irrigated trees in field conditions increases the feasibility of using this parameter in automatic irrigation systems.     

Hydraulic architecture and water flow in growing grass tillers (Festuca arundinacea Schreb.)

The water relations and hydraulic architecture of growing grass tillers (Festuca arundinacea Schreb.) are reported. Evaporative flux density, E (mmol s^?1 m^?2), of individual leaf blades was measured gravimetrically by covering or excision of entire leaf blades. Values of E were similar for mature and elongating leaf blades, averaging 2·4 mmol s^?1 m^?2. Measured axial hydraulic conductivity, K_h (mmol s^?1 mm MPa^?1), of excised leaf segments was three times lower than theoretical hydraulic conductivity (K_t) calculated using the Poiseuille equation and measurements of vessel number and diameter. K_t was corrected (K_t*) to account for the discrepancy between K_h and K_t and for immature xylem in the basal expanding region of elongating leaves. From base to tip of mature leaves the pattern of K_t* was bell‐shaped with a maximum near the sheath–blade joint (≈ 19 mmol s^?1 mm MPa^?1). In elongating leaves, immature xylem in the basal growing region led to a much lower K_t*. As the first metaxylem matured, K_t* increased by 10‐fold. The hydraulic conductances of the whole root system, (mmol s^?1 MPa^?1) and leaf blades, (mmol s^?1 MPa^?1) were measured by a vacuum induced water flow technique. and were linearly related to the leaf area downstream. Approximately 65% of the resistance to water flow within the plant resided in the leaf blade. An electric‐analogue computer model was used to calculate the leaf blade area‐specific radial hydraulic conductivity, (mmol s^?1 m^?2 MPa^?1), using , K_t* and water flux values. values decreased with leaf age, from 21·2 mmol s^?1 m^?2 MPa^?1 in rapidly elongating leaf to 7·2 mmol s^?1 m^?2 MPa^?1 in mature leaf. Comparison of and values showed that ≈ 90% of the resistance to water flow within the blades resided in the liquid extra‐vascular path. The same algorithm was then used to compute the xylem and extravascular water potential drop along the liquid water path in the plant under steady state conditions. Predicted and measured water potentials matched well. The hydraulic design of the mature leaf resulted in low and quite constant xylem water potential gradient (≈ 0·3 MPa m^?1) throughout the plant. Much of the water potential drop within mature leaves occurred within a tenth of millimetre in the blade, between the xylem vessels and the site of water evaporation within the mesophyll. In elongating leaves, the low K_t* in the basal growth zone dramatically increased the local xylem water potential gradient (≈ 2·0 MPa m^?1) there. In the leaf elongation zone the growth‐induced water potential difference was ≈ 0·2 MPa.     

Sap flow, gas exchange, and hydraulic conductance of young apricot trees growing under a shading net and different water supplies

The experiment was carried out in a research field near Murcia, Spain, over a 3-week period between September 26 and October 16, 2000. Sixteen trees were used in the experiment, eight of which were placed under a rectangular shading net, while the other eight were maintained in the open air. Trees were irrigated once per day and, after October 5th, water was witheld from eight trees (four shaded and four unprotected for 5 days). The leaf stomatal conductance and the photosynthesis rates were higher in the shaded trees than in the exposed plants, probably because the leaf water potential was lower in the unshaded plants. This higher leaf conductance partially compensated for the effect of low radiation on transpiration, and the reduction of daily sap flow registered in shaded trees was only around 10-20%. The net also affected trunk diameter changes, with the shaded trees showing lower values of maximum daily shrinkage. Soil water deficit and high radiation had a similar effect on plant water parameters, lowering leaf water potential, leaf stomatal conductance, and the photosynthesis rate. The effects of both conditions were accumulative and so the exposed water-stressed plants showed the lowest values of total hydraulic resistance and water use efficiency, while the shaded well-irrigated trees registered the highest values for both parameters. For this reason, we think that net shading could be extended to apricot culture in many areas in which irrigation water is scarce and insolation is high.     

Daily variations in water relations of apricot trees under different irrigation regimes

Mature apricot (Prunus armeniaca L. cv. Búlida) trees, growing under field conditions, were submitted to two drip irrigation treatments: a control (T1), irrigated to 100 % of seasonal crop evapotranspiration (ETc), and a continuous deficit (T2), irrigated to 50 % of the control throughout the year. The behaviour of leaf water potential and its components, leaf conductance and net photosynthesis were studied at three different times during the growing season, when they revealed a diurnal and seasonal pattern in response to water stress, evaporative demand of the atmosphere and leaf age. The deficit-irrigated trees showed, among other effects, a pronounced decrease in leaf water potential (ψ_w), decreased in leaf conductance (g_s) and no osmotic adjustment. For this reason, g_l and ψ_w can be considered good indicators of mature apricot tree water status and can therefore be used for irrigation scheduling.     

Estimation of hydraulic conductance within field-grown apricot using sap flow measurements

Using the heat pulse and other techniques, the hydraulic architecture of apricot trees was mapped out. The flows (overall flow, flow across the four main branches) and forces (water potential differences between xylem and leaves) measured allowed us to quantify hydraulic conductance of branches and of the root/soil resistance. The experiment was carried out in a commercial orchard of 11-year-old apricot trees (Prunus armeniaca L., cv. Búlida, on Real Fino apricot rootstock) during 1 week (October 27–November 3, 1998). Three representative trees with a cylindrical trunk divided into four main branches of different sizes, orientation and local microclimate were chosen for the experiment. Sap flow was measured throughout the experimental period. Twelve sets of heat-pulse probes were used, one for each main branch. The diurnal course of the environmental conditions, the fraction of the area irradiated and leaf water relations were also considered in each main branch. The relationships between leaf water potential, xylem water potential and transpiration were established for different branches and also for the total plant. Using the slopes of these regressions, total plant conductance, the hydraulic conductance of the stem and root pathway, the hydraulic conductance of the canopy and the hydraulic conductance of each branch were estimated. Our findings show that the root conductance and the canopy hydraulic conductance are similar in magnitude. Leaf hydraulic conductance per leaf area unit was similar for each of the four branch orientations, indicating that, while the light microclimate has a dominant influence on transpiration, in this case it had little effect on the hydraulic properties of the canopy.     

The effect of short-term flooding on the sap flow, gas exchange and hydraulic conductivity of young apricot trees

The effect of short-term flooding was examined in 2-year-old apricot trees (Prunus armeniaca cv. Búlida). Six apricot trees of similar appearance were submitted to two treatments: three were irrigated daily, while the others were flooded for a period of 50 h by submerging the pots in plastic water tanks. The trees were removed from the water, drained and then placed in the same conditions as the control plants. A decrease in transpiration in the flooded trees with respect to the control plants was evident. The daily pattern of soil O₂ concentration and plant hydraulic resistance followed a similar trend during the flooding. However, this relationship was not maintained throughout the experiment, since the O₂ values increased rapidly when the waterlogging ceased, while plant hydraulic resistance only recovered at the end of the experiment when the original root system, damaged by flooded conditions, was replaced with new roots. In flooded trees, the midday leaf water potential decreased progressively from the beginning of flooding, but gradually recovered when the waterlogging ceased. Leaf conductance values of treated plants were slow to recover, reaching values of the control plants 8 days after the leaf water potential had recovered. The close relationship observed during most of the experiment between the leaf water parameters, leaf conductance and plant hydraulic conductance indicate that hydraulic messages are likely to play a dominant role in co-ordinating the observed responses of the shoot.     

Distribution of hydraulic resistance in seedlings,sprouts and an adult tree ofPasania edulis Makino

Hydraulic resistance is an important factor in predicting water status. Hydraulic resistance of petiols, stems and branches, and roots was measured inPasania edulis Makino in order to compare the distribution of resistance between current seedlings, current stump sprouts and a 16 year old adult tree. Total resistance showed only minor variations despite large variations in plant size. This result is thought to be consistent with allometry between leaf mass and supportive organ mass, and with changes in permeability of conductive organs. Root resistance was low in sprouts and the adult tree due to their mature root systems. Current seedlings with undeveloped root systems had high root resistance. The proportion of petiol resistance in total resistance was high compared to the proportion of their conductive distance, and was thought to be a limiting factor of tree water status. The petiol resistance of the adult tree leaves was higher than for seedling and sprout leaves. From a comparison with the leaf water relation characteristics, the petiol resistance was thought to be provided as low values for intolerant leaves against water stress in order to compensate water inflow, and high values for tolerant leaves to regulate water inflow.     

Drought resistance of 2-year-old saplings of Mediterranean forest trees in the field: relations between water relations,hydraulics and productivity

Drought resistance was examined for 2-year-old saplings of Ceratonia siliqua L., Olea oleaster Hoffmgg. et Link., Quercus suber L. and Q. pubescens Willd. growing in the field in Sicily, with the aim of testing their possible use in the reforestation of degraded areas. To this purpose, leaf conductance to water vapour (g _L), transpiration rate (E _L), relative water content (RWC) and water potential (_L) were measured between pre-dawn and sunset, monthly from May to November. Parallel measurements of loss of hydraulic conductance of twigs of the current year (PLC) were made together with an estimate of whole-plant hydraulic conductance (K _PLANT) on the basis of the ratio of maximum E _L to (_PD-_MIN), where _PD is pre-dawn _L and _MIN is the minimum diurnal _L. C. siliqua saplings maintained high g _L throughout the study period with high RWC (over 90%) and _L. They grew rapidly and increased their foliage area (A _L) by over 60% from May to winter rest. This was accompanied by low twig PLC (about 20% in September) resulting in high K _PLANT all through the study period. In contrast, O. oleaster saplings underwent distinct dehydration in July, i.e. they showed stomatal closure due to a drop in RWC (to 75%) and _L (to the turgor-loss point). This was apparently due to twig cavitation, resulting in a strong decrease of K _PLANT. Plants, however, tolerated summer drought, and showed growth (A _L increased by 15% from May to November). Saplings of Q. suber and Q. pubescens were much more vulnerable to twig cavitation (PLC was 35–48% from June to November) than the other two species, and their average K _PLANT was lower. Saplings of Q. suber and Q. pubescens grew only during the wet spring period, and no new foliage was produced thereafter. Changes in twig hydraulic conductance played a dominant role in determining changes in K _PLANT in that the two variables were well correlated to each other (r=0.68 for P=0.001). We conclude that C. siliqua is an ideal candidate for reforestation of Sicilian degraded areas as is O. oleaster that, however, requires some additional water supply in the summer, at least during the early years after plantation. In wetter locations of Sicily, to an altitude between 0 and 500 m, Q. suber and Q. pubescens can be used for reforestation with expected higher competitiveness of the latter over the former species.     

Compensation heat-pulse measurements of sap flow for estimating transpiration in young lemon trees

Potted two-year-old lemon trees [Citrus limon (L.) Burm. f.], cv. Verna grafted on sour orange (C. aurantium L.) rootstock, growing in greenhouse, were subjected to drought for 33 d. Control plants were daily irrigated at field capacity. Values of sap flow (SF) were compared with transpiration (E) rates measured gravimetrically. The results underlined the robustness and high sensitivity of the compensation heat-pulse technique for estimating transpiration on a wide range of SF. Good direct correlations between E and SF rates on an instantaneous and daily basis were obtained in both treatments. On a daily basis, a common calibration curve can be used for both irrigation treatments. On an instantaneous basis, changes in SF were matches by similar changes in E in both treatments, although the relationships between these parameters presented different intercepts in each treatment. Sap flow rates were influenced by weather conditions in trees growing in non-limiting soil water conditions. This makes it possible to evaluate the significance of any sap flow measurement in relation to the reference value calculated for the vapour pressure deficit at the time the measurement was taken.This research was supported by Ministerio de Educacion y Ciencia (MEC), (CICYT/FEDER AGL2003-9387-C05-02 and AGL2004-0794-C03-02), and PETRI (PTR1995-0693-OP-02-01) grants to the authors. M.F. Ortuno was a recipient of a Program I3P research fellowship from CSIC.     

Drought resistance of Quercus pubescens as a function of root hydraulic conductance, xylem embolism and hydraulic architecture

Water relations, xylem embolism, root and shoot hydraulic conductance of both young plants in the field and potted seedlings of Quercus pubescens have been studied with the aim of investigating whether these variables may account for the well known adaptation of this oak species to arid habitats. Our data revealed that Q. pubescens is able to maintain high leaf relative water contents under water stress conditions. In fact, relative water contents measured in summer (July) did not differ from those recorded in April. This was apparently achieved by compensating water loss by an equal amount of water uptake. Such a drought avoidance strategy was made possible by the recorded high hydraulic efficiency of stems and roots under water stress. In fact, root hydraulic conductance of field-grown plants was maintained high in summer when the percentage loss of hydraulic conductance of stems was lowest. The hydraulic architecture of young plants of Q. pubescens measured in terms of partitioning of hydraulic resistances along the water pathway revealed that the highest hydraulic resistance was located in stems of the current year's growth. This hydraulic architecture is interpreted as consistent with the adaptation of Q. pubescens to arid habitats as a consequence of the recorded seasonal changes in water relation parameters as well as in root and stem hydraulics.     

Water status indicators of lemon trees in response to flooding and recovery

Potted 2-year-old lemon trees [Citrus limon (L.) Burm. fil, cv. Verna] grafted on sour orange (C. aurantium L.) rootstock were subjected to flooding for 3 d. Control plants were irrigated daily to field capacity. Continuously (sap flow, trunk diameter fluctuations) and discretely (predawn and midday leaf water potential, leaf conductance) measured plant-based water status indicators were compared. The sensitivity of the maximum daily trunk shrinkage signal intensity to flooding and its behaviour during the recovery period demonstrated that this indicator is more feasible than the others for use in automatic irrigation. The responses to flooding of continuously and discretely measured plant-based water status indicators were very similar to those observed in response to drought stress indicating that it necessary to use soil water measurement automatic sensors to detect the cause of the stress. The results underlined the robustness of the compensation heat-pulse technique for estimating instantaneous and daily transpiration rates on flooding stress and recovery.     

Leaf water potential-leaf water deficit relationship for ten species of a semiarid grassland community

The relationship between water potential and relative water content (water content in percentage of full hydration) is a characteristic of plant tissues, that may vary with environmental conditions. It is used here to compare leaf water relations of ten species coexisting in a semiarid grassland community (Festucetum vaginatae danubiale) in Hungary. Three groups of species can be distinguished. In two of these leaf water potential changes only moderately with decreasing leaf water content. These are either short-lived, drought escaping spring plants relying on seasonally favourable water supply (group 1) or xerophytes with very deep root system having access to permanent water resources (group 2, only one species studied here). Xerophytes with moderately deep roots (group 3) display a rapid drop of leaf water potential with increasing leaf water deficit. This generates a steep water potential gradient in the soil-plant continuum that in turn enhances water uptake by roots. There is a positive correlation between the rate of water potential decline and degree of sclerophylly (proportion of dry material in the water-saturated leaf), and both variables show seasonal change in perennial species.     

Hydraulic Properties of a Mangrove Avicennia Germinans as Affected by NaCl

Water transport was assessed in seedlings of the mangrove Avicennia germinans L. grown at 171 and 684 mol m^–3 NaCl. Leaf specific conductivity declined by 25 % at high salinity. This was related to low specific conductivity, because Huber values remained similar. Leaves of A. germinans featured low internal conductance to water transport. This was lowered further under high salinity. Water transport constrains imposed by whole shoot and leaf blade at high salinity were balanced by stomatal regulation of water loss, which possibly maintain stem water potentials above embolisms levels.     

Relationship between transpiration and water potential in grafted scions of Picea

Total water and osmotic potential, turgor pressure and transpiration rate were measured on scions of Picea pungens (Englemann) during union development. In controlled environments, declines in water potential were correlated with lower transpiration rates to about −2.0 MPa. Water potentials below −2.0 MPa resulted in graft failure and were associated with sharply increased transpiration rates. Bulk turgor pressures remained high in the needles during this period of declining water potential and increasing transpiration. Transpiration rates of successful and unsuccessful greenhouse grafts were not significantly different during union development. Transpiration rates of these grafts were highest around dawn, then declined throughout the day only to increase again after sunset. High bulk needle turgor values (1.3 MPa), maintained by osmotic adjustment, may prevent stomatal closure of Picea scions at water potentials below −2.0 MPa.     

Long-term transpiration change with rainfall decline in a Mediterranean Quercus ilex forest

In the Mediterranean basin, precipitation is expected to decline as a consequence of climate change. The response of a Quercus ilex forest in southern France to such a decline in water availability was studied using a 4-year throughfall exclusion experiment. Seasonal courses of sap flow and leaf water potential were obtained from 2004 to 2007 and used to characterize tree water relations in a control and a dry treatment. The experiment reduced the average precipitation input to the soil by 29%, and resulted in a 23% reduction in annual transpiration. Soil water potential was significantly lower in the dry treatment only during summer drought, but transpiration was reduced all year round even during well-watered periods. Despite a tight stomatal control over transpiration, whole-tree hydraulic conductance was found to be lower in the trees growing in the driest conditions. This reduction in water transport capacity was observed jointly with a reduction in leaf transpiring area. Canopy leaf area decreased by 18% in the dry treatment as a consequence of the throughfall exclusion, which was found to validate the ecohydrological equilibrium theory.     

Effect of water deficits on transpiration, photosynthesis and leaf conductance in cassava

Transpiration, net photosynthesis and leaf conductance decreased when leaf water potential dropped below -0.30 MPa. Both transpiration and net photosynthesis rates were considerably reduced before the leaves were visibly wilted at -0.95 MPa. Consequently, visual symptoms are unlikely to provide a useful index for characterizing water deficits in cassava ( Manihot esculenta Crantz cv. Llanera). Decreases in net photosynthesis closely followed decreases in transpiration and this suggests that stomatal closure controls both processes.     

Phase and amplitude relations between transpiration, water potential and stem shrinkage

Abstract An electrical analogue describing the phase and amplitude relations between transpiration, water potential and stem shrinkage for trees was developed. Observations of shrinking and swelling at various heights up a Pinus radiata tree were obtained over several weeks in summer and autumn. The relative amplitude in shrinkage increased by a factor of two up the stem, but phase lags were small. The data obtained were used in conjunction with the electrical analogue of the How pathway to obtain an estimate of the relative magnitude of the capacitance of the living bark and the sapwood, and to predict phase lags between transpiration and leaf water potential, and lags in transpirational flux up the stem. The results suggest that if water potentials recover by dawn, phase lags in water potential down tree stems arc small and that the exchange between water stored in the sapwood and the transpiration stream is irreversible over the diurnal time scale.