To be useful for indicating plant water needs, any measure of plant stress should be closely related to some of the known short- and medium-term plant stress responses, such as stomatal closure and reduced rates of expansive growth. Midday stem water potential (SWP) has proven to be a useful index of stress in a number of fruit trees. Day-to-day fluctuations in SWP under well-irrigated conditions is well-correlated to midday vapor pressure deficit, and hence can be used to predict a non-stressed baseline. Measurement of SWP helped to explain the results of a 3-year deficit irrigation study in mature prunes, which showed that deficit irrigation could have either positive or negative impacts on tree productivity, depending on soil conditions. Mild to moderate water stress was economically beneficial. In cherry, SWP was correlated to both leaf stomatal conductance and rates of shoot growth, with shoot growth essentially stopping once SWP dropped to between –1.5 to –1.7 MPa. In pear, increased fruit size, decreased fruit soluble solids, and increased green color were all associated with increases in SWP.
Kenneth A. Shackel
Rémy E. Milad and Kenneth A. Shackel
Irrigation of previously water-stressed French prune trees is known to induce fruit end cracking. The relationships between end cracking, water relations, and mechanical properties of the skin of French prune were studied as a function of irrigation regimes under field conditions. Water stress resulted in the accumulation of solutes in the fruit of nonirrigated trees. A gradient in osmotic potential (ΨS) existed along the vertical axis of fruit from all treatments; ΨS was always lower at the stylar than stem end. Irrigation of previously water-stressed trees (irrigated-dry treatment) resulted in ΨS gradients exceeding those of all other treatments. Moreover, estimated turgor (ΨP) at the stylar end of the fruit increased 2-fold within 24 hours after irrigation. These changes were accompanied by the onset of fruit end cracking, and neither the well-watered controls nor the continuously droughted fruit exhibited such changes. During the 24 hours following irrigation, the overall ΨS of irrigated-dry treatment fruit was diluted by the same amount as the calculated increase in fruit volume. However, during the same period, ΨS at the stem end of the fruit showed more dilution than expected, and ΨS at the stylar end of the fruit concentrated, indicating a redistribution of solutes. There were no differences in skin mechanical properties along the fruit vertical axis and, hence, this could not have accounted for the observed changes in ΨS and ΨP. Thus, when previously stressed French prune trees were irrigated, the overall recovery in water potential (Ψ) and the subsequent movement of solutes to the stylar end of the fruit resulted in apparently excessive turgors in this region and hence the observed pattern of end cracking.
Bruce D. Lampinen and Kenneth A. Shackel
Two levels of deficit irrigation were applied to mature French prune trees based on gradually declining midday stem water potentials decreasing to –1.5 MPa (moderate stress) and –2.5 MPa (severe stress) by harvest. The moderate and severe stress treatments resulted in 32% and 51% water savings, respectively, compared to the fully irrigated control. The average photosynthetic rate and dry fruit yield for the moderate stress treatment were not significantly lower than those for the fully irrigated control. The severe stress treatment had significantly lower photosynthetic rates from late morning until sunset on most days. However, this lower photosynthetic rate did not result in significantly lower dry fruit yields. There were no significant differences in light interception as measured diurnally or over the course of the season in the first year of the study. However, light interception and photosynthetic differences might be expected to be more pronounced in subsequent years, due to carryover effects.
Sunitha H. Gurusinghe and Kenneth A. Shackel
Commercial harvesting of almonds [Prunus dulcis (Mill.) Webb.] with mechanical shakers is economical, but may also cause severe damage to the cambial zone, leading to the establishment of a deadly cankerous disease, ceratocystis. Irrigation is often cut off for some period of time before harvest, anticipating a reduction in the damage as a result of an increased strength of adhesion within the cambial zone between the bark and the wood. Mechanical failure can occur in the cambial zone proper and in the differentiating cells on either side of the cambial zone. In this study the shear force per shear area in the cambial zone (cambial strength) was measured to represent mechanical failure due to shaker damage. Cambial strength of branches with intact bark tissues could be reversibly influenced by the level of tissue hydration, with high cambial strength associated with increased tissue hydration, presumably a result of an increase in the turgor of cambial zone cells. Methods of measuring cambial strength were developed for branches and trunks to avoid the effects of reversible turgor changes. Irrigation regimes imposed before harvest had a substantial and progressive influence on tree stem water potential, stomatal conductance, and growth rate of almond trees. Statistically significant levels of within seasonal differences in rate of growth, stomatal conductance and tree water deficits found under different irrigation regimes did not show any correlation with the within seasonal fluctuations in cambial strength. Cambial strength always showed a similar pattern and a similar magnitude of seasonal increase from spring to summer (as previous authors have reported), superimposed over the within seasonal fluctuations, despite significant differences in tree water stress. Therefore, this study suggests that irrigation cut off may be an ineffective practice for the purpose of increasing cambial strength.
Sunitha H. Gurusinghe and Kenneth A. Shackel
Mechanized shaker harvesting of large acreages of almond [Prunus dulcis (Mill.) Webb.] trees leads to economical use of labor, general cost reduction, and speed of harvest. However, shaking can separate the bark (all tissues external to the vascular cambium) from the wood (all tissues internal to the vascular cambium) and exposes the vascular cambial zone to infection by the fungus Ceratocystis fimbriata Ell. & Halst. leading to mallet wound canker and loss of productivity. Treatment of ethephon caused a significant increase in the cambial strength of both branches and trunks, presumably by ethylene-induced anatomical and biochemical changes in the cambial zone. An increase in the ratio of total tangential area of the groups of ray initials to fusiform initials and the thickness of ray initial cell walls was observed as a response to ethephon treatment. Spraying an ethephon solution (500 μl-liter-1) to the runoff point on almond trunks caused significant increases in the cambial strength; ethephon, therefore, may be useful in reducing bark injury during shaker harvesting.
Rémy E Milad and Kenneth A Shackel
End cracking of French prune fruits occurs when previously water stressed trees are irrigated during early July. Fruit phloem, xylem and transpiration flows (P, X and T, respectively) were measured diurnally during 72 h periods in mid June, early July and mid July (before, during and after the crack-susceptible period). Midway through each 72 h period, the previously stressed trees were irrigated. In mid June, X was larger than P, whereas P was larger than X during early July. In mid July, P and X were similar. In early July, the period preceding irrigation was characterized by an ourflow of phloem sap during the day and phloem inflow during the night. After irrigation, larger phloem inflows were observed and no phloem outflow occurred. Fruit transpiration rates were highly correlated with VPD. They exhibited a gradual decrease during the season, reaching minimum values during early July, before increasing again. The sum of P and X was virtually identical for the three periods i.e. stronger P's compensated for weaker X's and vice versa. Our results suggest that properties intrinsic to the fruit play the primary role in modulating water and photosynthate movements between the tree and the fruit. The possible role of these properties on fruit growth and cracking will be examined.
Juan C. Díaz-Pérez, Kenneth A. Shackel, and Ellen G. Sutter
Little is known about the physiological changes that occur during acclimatization and how these changes influence plant survival and growth in the new environment. In particular, it is unclear to what extent in vitro-formed roots are functional in water uptake, particularly when the plantlet is exposed to conditions of increasing evaporative demand. Tissue-cultured shoots and plantlets (shoots with roots) were acclimatized by exposing them to a linear reduction in relative humidity (RH) from 99 % to 75%over 4 days. When conductance was measured at 95% RH (21 C), in vitro shoots and plantlets showed a very high initial conductance, followed by a gradual decline, reaching steady state in 12 hours. Acclimatized shoots and plantlets had a 50% lower initial conductance compared to nonacclimatized ones, and reached steady state in 4 hours. The reduction in conductance as a result of acclimatization most likely contributes to a reduced transpiration under conditions of increased evaporative demand. Roots formed in vitro were associated with a higher plant water status, suggesting that these roots were functional in water uptake. Relative water content of the shoot was positively correlated with leaf conductance and net photosynthesis. We suggest that tissue-cultured plantlets behave as hydraulically integrated units, in which there must be a coordination between control of water loss by the shoot and uptake of water by the root to maintain a favorable plant water balance. Our results also indicate that methods that use excised shoots or leaves to determine transpiration gravimetrically may not accurately represent the stomatal water loss characteristics of tissue-cultured plants.
Kenneth A. Shackel, H. Ahmadi, C. Greve, and J. Labavitch
The pressure microprobe was used to measure cell turgor (ψp) in tomato pericarp tissue, and also to sample vacuolar fluid for the measurement of cell osmotic potential (ψs) in a nanoleter freezing point osmometer. In fresh tissue, cell ψs agreed well with the ψs of frozen-thawed whole tissue measured with a vapor pressure osmometer. Under a wide range of ripeness conditions however, and for both intact fruit and discs of fruit tissue, fruit cell turgor was consistently lower than expected, based on the values of cell ψs. When tissue discs were hydrated in aerated distilled water, disc fresh weight increased substantially (20 - 50+%), and both cell turgor and tissue ψs increased. Cell ψs however, remained relatively constant. These and other observations suggest that the turgor increase during hydration was largely due to losses of solute from the apoplastic space, partly by direct losses from the tissue, and partly by cell solute accumulation.
Soo-Hyung Kim, Kenneth A. Shackel, and J. Heinrich Lieth
Shoot-bending has become a standard cultural practice in cut-flower rose (Rosa hybrida L.) production. Physiological effects of shoot-bending on leaf net photosynthesis (A), stomatal conductance (gs), transpiration rate (E), and stem water potential (ψ) were investigated for rose plants. With saturating light conditions, shoot-bending decreased rates of A, gs, and E in comparison with the rates prior to shoot-bending. A, gs, and E of bent shoots were significantly lower than those of the control shoots that were not bent. The differences in A between bent and control shoots decreased over time, disappearing within 3 weeks after bending. Bent shoots exhibited reduced ψ. Leaves projecting upward on a bent stem were found to have higher A, gs, and E than those projecting downward. This was probably due to the destruction of xylem vessels serving the leaves attached to the lower side (compression side) of the bent stem. Our results support the hypothesis that hydraulic conductivity is reduced in bent shoots probably due to disturbed xylem tissues, and that reduced photosynthetic rates of bent shoots are a function of water status.
Kenneth A. Shackel, R. Scott Johnson, Charles K. Medawar, and Claude J. Phene
The heat balance method was used to estimate transpirational sap flow through 60- to 75-mm-diameter stems (trunks) of 3-year-old peach [Prunus persica (L.) Batsch. cv. O'Henry] trees under field conditions. On rare occasions, heat balance estimates agreed well with independent lysimetric measurements, but on most occasions, heat balance estimates of sap flow were unrealistic in both direction and magnitude. In some cases, the errors in sap flow approached two orders of magnitude and were always the result of a calculation involving division by a very small and sometimes negative temperature differential between the stem surface temperature above and below the gauge heater. The occurrence of negative temperature differentials under positive transpiration conditions may be inconsistent with a fundamental assumption in the heat balance model, namely that temperature differentials are solely a consequence of the dissipation of energy supplied to the gauge heater. In the absence of heating power applied to the gauge, temperature differentials exceeding - 1C were correlated with the rate of change in stem temperature, indicating that ambient conditions themselves can impose a bias in gauge signals and, hence, influence gauge accuracy. Our results suggest that the effect of ambient conditions on gauge signals should be critically evaluated before considering heat balance estimates of sap flow as reliable under any given conditions.