Seedlings of pecan [Carya illinoensis (Wangenh.) C. Koch] grown in perlite culture were treated with N and S in a 5 × 5 factorial in a randomized complete block design to determine the effect of N, S, and N × S interaction on vegetative growth and photosynthesis. Nitrogen and S deficiency symptoms occurred when leaf N and S were < 25 and 1.4 mg·g-1 dry weight, respectively. Photosynthesis was reduced when combined leaf N and S exceeded 35 and 3.7 mg·g-1 dry weight, respectively; growth was reduced when leaf N and S were > 34 and 3.7 mg·g-1 dry weight, respectively. Photosynthesis and growth increased with N supply, but depended on leaf N: S ratio. In plants without visible N or S deficiency, a N: S ratio of ≈9. is proposed to be near the optimum for maximum growth. Comparison of leaf N, S, and the N: S ratio with similar analyses in selected orchards suggests that pecan productivity will increase from S application under field conditions. We conclude that the interaction” of N and S imposes stringent controls on leaf N and S, photosynthesis, and growth.
Hening Hu and Darrell Sparks
Hening Hu and Darrell Sparks
Leaves of `Stuart' pecan [Carya illinoensis (Wangenh.) C. Koch] with various levels of Zn deficiency were analyzed for physiological indicators of leaf vigor. Leaf chlorophyll content, stomatal conductance, and net photosynthesis were adversely affected by Zn deficiency. In leaves with severe Zn deficiency, each of these indicators increased 3- to 5-fold with a doubling of leaf Zn concentration, but stabilized as leaf Zn approached the sufficiency range (14 μg·g-1). High intercellular CO2 associated with low net photosynthesis indicates that stomatal aperture was not the cause of the reduction of net photosynthesis under Zn deficiency.
Hening Hu and Darrell Sparks
The effect of Zn deficiency on reproductive growth of `Stuart' pecan [Carya illinoensis (Wangenh.) C. Koch] was studied. At the most severe Zn-deficiency level, shoots were rosetted and produced neither. staminate nor pistillate inflorescences. At less severe Zn-deficiency levels, catkin length and weight decreased as Zn concentration in the leaf decreased. The number of fruits produced per shoot was reduced by Zn deficiency. Even though fruit abortion was not affected by Zn status of the shoot, fruit death and drying in situ increased with increasing Zn deficiency. Zinc deficiency dramatically suppressed fruit development and resulted in delayed and staggered shuck dehiscence.
Hening Hu and Gary A. Couvillon
The activities of catalase and of glucose-6-phosphate dehydrogenase (G6PDH) and 6-phosphogluconate dehydrogenase (6PGDH), the two key enzymes in the pentose phosphate pathway (ppp), were measured in the seeds of Prunus persica (L.) Batsch var. nectarina Maxim `Nectarine 7'. The seeds were subjected to three imbibition treatments: 1) continuous 24C; 2) continuous 4C; and 3) application of thiourea (TU)/gibberellic acid (GA) at various concentrations to seed held at 24C then subsequently chilled at 4C. Treatments of continuous 24 or 4C indicated that catalase, G6PDH, and 6PGDH exhibited significant activity increases only when the seeds obtained germination potential, which occurred in the seeds chilled for 7 weeks at 4C. Seeds held at 24C did not germinate and showed little change with time in G6PDH and 6PGDH activity. There was only a slight increase in catalase activity beginning 3 weeks following treatment initiation and a decrease in activity following 13 weeks of treatment. Thiourea treatment resulted in an inhibition of catalase activity and a stimulation of G6PDH, but had no effect on 6PGDH activity. However, no correlation between enzymic activity and seed germination was found. The results strongly questioned the role of the ppp and catalase activity in dormancy control as previously hypothesized.
Hening Hu, Darrell Sparks, and John J. Evans
Greenhouse-grown pecan [Carya illinoensis (Wangenh.) C. Koch] seedlings were treated with S as SO4 -2 (0 to 4 mm) to determine the effect of S on its vegetative growth, chlorophyll concentration, nutrition, and free and protein amino acid content. Sulfur deficiency symptoms occurred when leaf S was ≈1.5 mg·g-1 dry weight or less. Chlorophyll concentration and growth increased curvilinearly with leaf S and had not peaked when leaf S was 2.7 mg·g-1 dry weight. The N : S ratio in the leaf and leaf S were equally reliable indicators of the S status of the plant, but the optimum ratio was less than the value of 15 found in other plant species. Calcium and Mg decreased with S application, suggesting a direct SO4 -2 or NH4 + effect. The concentration of all other elements determined was elevated, in one or more plant parts, when S deficiency symptoms were visible. The most conspicuous effect of S deficiency on N metabolism was a greatly expanded free amino acid pool, mainly arginine. The concentration of free amino acids decreased exponentially with leaf S. Conversely, protein amino acid was inhibited by S deficiency and increased with leaf S concentration or, perhaps more specifically, with methiouine.
Rawia El-Motaium, Hening Hu, and Patrick H. Brown
The influence of B and salinity [3 Na2SO4 : 1 CaCl2, (molar ratio)] on B toxicity and the accumulation of B, sodium, and SO4 in six Prunus rootstocks was evaluated. High salinity reduced B uptake, stem B concentrations, and the severity of toxicity symptoms in five of the six rootstocks. Forward and backward stepwise regression analyses suggested that stem death (the major symptom observed) was related solely to the accumulation of B in the stem tissue in all rootstocks. The accumulation of B and the expression of toxicity symptoms increased with time and affected rootstock survival. No symptoms of B toxicity were observed in leaf tissue. The Prunus rootstocks studied differed greatly in stem B accumulation and sensitivity to B. The plum rootstock `Myrobalan' and the peach-almond hybrid `Bright's Hybrid' were the most tolerant of high B and salinity, whereas the peach rootstock `Nemared' was very sensitive to high B and salinity. In all rootstocks, adding B to the growth medium greatly depressed stem SO4 concentrations. In every rootstock except `Nemared' peach, adding salt significantly depressed tissue B concentrations. A strong negative correlation between tissue SO4 and B was observed. Grafting experiments, in which almond was grafted onto `Nemared' peach or `Bright's Hybrid', demonstrated the ability of rootstocks to influence B accumulation and scion survival.
Patrick H. Brown, Hening Hu, and Rawia El-Mntaium
Excessive soil and water boron (B) occur widely in California, often in conjunction with high soil salinity. Descriptions of the symptoms of B toxicity and quantification of its impact on Prunus species are not available. In these experiments we describe the impact of high B and saline conditions on uptake, distribution and growth depression in almond, peach, plum and peach/almond hybrid rootstocks
A series of experiments are described that indicate an important additive effect of B on sensitivity of Prunus species to salinity. Boron concentrations in excess of lppm in the irrigation solution, significantly impair plant growth under moderate (non-limiting) salinity conditions and lead to plant death at higher salinity levels. Symptoms of B toxicity in Prunus include stem necrosis and vascular occlusion. Unlike most other species, B does not appear to accumulate in the leaf margins and leaf symptoms are generally not observed. Differences in sensitivity of a range of Prunus species to B toxicity are described.
Initial results suggest that differences in rootstock sensitivity to B and salinity are the result of differential uptkake and partitioning of B, Na and Cl within the plant.
Patrick H. Brown, Hening Hu, and Warren G. Roberts
The phloem mobility of boron (B) in plants varies dramatically among species. Variations in phloem B mobility occur as a consequence of the presence of sugar alcohols (polyols) in some species but not in others, and these differences in phloem B mobility profoundly affect the expression of B toxicity symptoms. Twenty-four species including common ornamental species varying in sugar alcohol content, were selected to test their response to B toxicity. Species that do not produce sugar alcohols exhibited previously described B toxicity symptoms that include accumulation of high concentrations of B in, and burning of, the tip and margin of old leaves. In the sugar-alcohol-producing species these symptoms were absent, and B toxicity was expressed as meristematic dieback and an accumulation of B in apical tissues. These symptoms have not previously been associated with B toxicity in these species and hence may have been frequently misdiagnosed.
Slavko Perica, Nacer Bellaloui, Carl Greve, Hening Hu, and Patrick H. Brown
Boron (B) remobilization, mannitol and glucose concentrations, and the effect of B application on changes in soluble carbohydrates were investigated in various organs of bearing `Manzanillo' olive trees (Olea europaea L. `Manzanillo'). Following foliar 10B application to leaves of various ages, there was significant 10B export out of the treated leaves, and significant 10B enrichment in nontreated adjacent organs, including inflorescences and fruit. Results demonstrated that B can be remobilized from leaves of various ages, and that foliar-applied B is phloem mobile in olive. Soluble carbohydrate analysis determined that mannitol and glucose are the predominant sugars in all organs analyzed and that the mannitol concentration in the leaves is adequate to account for all B transport. This is consistent with observations in other species, where the presence of mannitol is known to facilitate phloem B transport through formation of a mannitol-B complex. Previous reports have indicated that B application can alter carbohydrate metabolism. In the present study, foliar B application significantly suppressed glucose concentration in the leaf petioles of all ages and increased mannitol in petioles of the current-year-developed leaves.
Slavko Perica, Patrick H. Brown, Joseph H. Connell, Agnes M.S. Nyomora, Christos Dordas, Hening Hu, and James Stangoulis
A 2-year field study was conducted to determine if foliar B applications prior to flowering increased fruit set in olive (Olea europaea L.) cv. Manzanillo. Boron solutions were applied (935 L·ha-1) at four concentrations (0, 246, 491, and 737 mg·L-1) to trees exhibiting no vegetative symptoms of B deficiency. Foliar B application increased both the percentage of perfect flowers and fruit set, but no effect on pollen germination was observed in either year. The increase in fruit set was not accompanied by a reduction in fruit size. The beneficial effects of foliar B application varied between years and were greater when fruit set was low. The results obtained here are in agreement with those observed in other tree species, in which foliar B applications made immediately prior to flowering or during the period of floral bud initiation significantly increased fruit set and yield. The physiological basis for this effect, however, remains unclear.