Changes in Photosynthesis and Fruit Characteristics in Olive in Response to Assimilate Availability

The source level in the olive cultivar Leccino was varied by girdling at different stages of fruit growth. Afterwards, the effects on gas exchange, fruit growth, and ripening and blooming were studied. Girdling during fruit growth did not significantly influence net photosynthetic rate (P _N) except in the last phase of fruit growth when the P _N was reduced. In the girdled branch, P _N began to decrease at the onset of starch accumulation because fruit growth ceased. In mid-November stomatal conductance (g _s) and transpiration rate (E) were also reduced by girdling, whereas sub-stomatal CO₂ concentration (C _i) increased in leaves from the girdled branches. The total chlorophyll content (Chl) tended to decrease in parallel with the reduced P _N. Girdling did not substantially influence the leaf and shoot water contents. The large availability of assimilates seems to cause an earlier fruit ripening. In general, girdling increased fruit dry mass. Healing before the time when the majority of pulp growth occurs reduced the effect of girdling. June girdling increased the pit dry mass. Girdling at the beginning of August and September, compared to the control, increased the pulp dry mass, but the pit dry mass did not differ with respect to the control. The percentage of oil in the fruit, on a dry mass basis, increased with August and September girdlings, but the percentage of oil in the pulp did not change. Girdling reduced shoot growth, but the internode length was unchanged. Girdling slightly stimulated differentiation of flower buds.     

Effect of Fruiting on Leaf Gas Exchange in Olive (Olea Europaea L.)

The effect on traits of photosynthesis and water relations of assimilate demand was studied in olive tree that has strong alternate bearing. The diurnal and seasonal leaf gas exchanges, area dry mass, and saccharide and chlorophyll (Chl) contents were measured by comparing shoots with fruit of "on-trees" (heavy fruit load) with shoots without fruit on both "on-trees" and "off-trees" (light fruit load). In spite of large seasonal and diurnal differences, leaf net photosynthetic rate (P _N), stomatal conductance (g _s), sub-stomatal CO₂ concentration (C ₁), transpiration rate (E), and respiration rate (R _D) were not significantly influenced by fruit load or by the presence or absence of fruit on the shoot. An only exception was at the beginning of July when the one-year-old leaves on shoots with fruit had slightly higher P _N and E than leaves on shoots without fruit. Water content, Chl and saccharide contents, and area dry mass of the leaf were not substantially influenced by the presence/absence of fruit on the shoot or fruit load. Hence the sink demand, associated with fruit growth, did not improve leaf photosynthetic efficiency in olive. This revised version was published online in August 2006 with corrections to the Cover Date.     

Fruit Set, Abscission and Dry Matter Accumulation on Girdled Branches of Macadamia

The patterns of fruit growth, abscission and dry matter accumulationwere determined for ungirdled macadamia branches and comparedwith the patterns on girdled branches. The relationships betweenfruit set, fruit drop, and the number of available leaves werealso assessed. Approximately 50 leaves were required to supportdevelopment of each fruit on girdled branches, so that finalfruit numbers on girdled branches were higher or lower thanon ungirdled controls, depending on leaf number. High leaf numbersdid not cause any increase in fruit size on girdled branches,but low leaf numbers resulted in fruit volumes up to 23% lowerthan the controls. The number of fruits set on ungirdled brancheswas independent of the number of leaves on the branch. Defoliationof ungirdled branches had no effect on the number of fruitsset, with fruit set on these branches apparently being supportedalmost entirely from carbohydrates obtained elsewhere in thetree. Three phases of fruit drop were observed, with maximaat 2, 6-7, and 10 weeks post-anthesis, separated by phases ofrelatively low fruit drop at 4 and 8 weeks post-anthesis. Themaximum dry weight increase of the crop occurred between 10and 20 weeks post-anthesis, when there was very little fruitdrop. Therefore, although limited availability of assimilatesmay influence fruit retention in macadamia, there is no simplerelationship between structural carbon demand and fruit drop.Copyright1994, 1999 Academic Press Macadamia integrifolia Maiden and Betche, Macadamia tetraphylla L. A. S. Johnson, Proteaceae, macadamia, fruit set, fruit drop, girdling     

Effects of shoot girdling on photosynthetic capacity, leaf carbohydrate, and bud abscission in pistachio (Pistacia vera L.)

The effects of shoot girdling on stomatal conductance (g _s), leaf photosynthesis (P _N), concentrations of carbohydrates, nitrogen and chlorophyll (Chl) in leaves, areal leaf mass (ALM), the diameter and length of shoots, and bud abscission in pistachio were investigated. Girdling individual shoots at the base of the current year’s shoot (girdle I), separating inflorescent buds on the terminal current year’s shoot from the developing fruits on the previous year’s shoot, reduced inflorescent bud abscission by 70% in comparison to nongirdled controls. Girdle I significantly reduced concentrations of nitrogen in leaves but increased those of nonstructural carbohydrates particularly of starch. Shoot diameter increased by 13.1% and 26.4% at 33 and 81 days after girdling (DAG), respectively, compared to 1% and 3.4% in the control, respectively. Both the leaf dry mass/fresh mass ratio and ALM were increased significantly by girdle I from 12 DAG. The concentrations of Chl a, Chl b, Chl (a+b), as well as the ratio of Chl a/b, all decreased with girdle I. The greatest negative effect of girdle I was on g _s and P _N. P _N was reduced by 55% of its initial value and was 44% less than in the control leaves at 10 DAG, and fell to approximately 30% that of the control from 21 DAG. In contrast, girdling at the base of one-year-old shoots (girdle II), thus not separating fruits from the inflorescent buds, did not significantly affect g _s or P _N. The effect of girdling on P _N and the possible factors that are involved in the reduction of photosynthesis in pistachio are discussed.     

Net CO(2) Assimilation Rate of Grapevine Leaves in Response to Trunk Girdling and Gibberellic Acid Application

Net CO₂ assimilation rate (A), stomatal conductance (g_s), and weight per unit leaf area (W) were determined on Thompson Seedless grapevines grown in the field. Treatments included fruit set applications of gibberellic acid (40 milligrams gibberellic acid (GA₃) per liter) to vines, shoots and clusters, alone and in combination with trunk girdling. Leaf A and g_s were measured prior to and 3, 6, and 13 days after fruit set. Weight per unit leaf area was determined on leaves collected subsequent to gas exchange measurements. Leaf A of girdled vines was reduced approximately 30% when compared to the control 13 days after treatment. The reduction in A due to girdling was not as great when vines were sprayed with GA₃. GA₃ sprays alone had no significant effect on A. Stomatal conductance was reduced by girdling 13 days after treatment. Weight per unit leaf area was 17% greater for trunk girdled vines when compared to the controls. Results indicate GA₃ affected net CO₂ assimilation rate only on girdled vines, a treatment which increased weight per unit leaf area.     

Impact of phloem girdling on leaf gas exchange and hydraulic conductance in hybrid aspen

We investigated phloem-xylem interactions in relation to leaf hydraulic capacity in hybrid aspen (Populus tremula L. × P. tremuloides Michx.) by using phloem girdling method. Removal of bark tissues (phloem girdling) at the branch base resulted in a substantial decline in stomatal conductance (g_S), net photosynthetic rate (P_N), and leaf hydraulic efficiency, and in increase of leaf water potential (Ψ_L). Although gS declined more than P_N (83 versus 78 %), the ratio of intercellular to ambient CO₂ concentrations (c_i/c_a) increased from 0.67 to 0.87 in three days after girdling. Girdling induced a decrease in leaf hydraulic conductance (K_L) on average by 43 % (P = 0.006). The changes in gS and leaf conductance to water vapour were co-ordinated with K_L only in girdled branches whereas intrinsic water-use efficiency was invariant to K_L. The declines in K_L with girdling were not accompanied by changes in potassium ion concentration ([K⁺]), electrical conductivity, or pH of xylem sap. The results suggest that phloem girdling at the branch base does not influence the recirculation of ions between the phloem and xylem in hybrid aspen and the decrease of K_L in response to the manipulation is not related to changes in [K⁺] and total ionic content of xylem sap.     

Effect of leaf-to-fruit ratio on leaf nitrogen content and net photosynthesis in girdled branches of Mangifera indica L.

Variations in leaf nitrogen concentration (N_m), mass-to-area ratio (M_a), amount of leaf nitrogen per unit leaf area (N_a), and non-structural carbohydrates were measured in well-lit girdled branches of 11-year-old mango trees that were experiencing similar conditions of irradiance and gap fraction. The influence of source–sink relationships was studied by testing three levels of leaf-to-fruit ratio: 40, 70 and 150, during the period of linear fruit growth. N_a was negatively correlated to the leaf-to-fruit ratio. Differences in N_a were reflected in differences in net photosynthetic assimilation, A_net, although they could not fully account for them. All differences in N_a resulted exclusively from differences in N_m, not M_a. Starch and total non-structural carbohydrates accumulated in the leaves as the result of higher leaf-to-fruit ratio, which suggests that the leaf carbohydrate status may play a role in photosynthetic acclimation to fruit load in mango. These observations complement previous findings where photosynthetic acclimation to light was found to be driven by changes in M_a, while N_m remained almost constant over a large range of gap fractions. Observations about the effect of fruit load were also in contrast with previous observations, since no evidence was found that leaf carbohydrate status played any role in photosynthetic acclimation to light. This study demonstrates that acclimation to changing source–sink relationships does not follow the same pattern as acclimation to progressive shading; but these observations do suggest that there may be different mechanisms by which leaves acclimatise to changing conditions.     

Photosynthesis,Transpiration, and Water Use Efficiency of Vegetative and Reproductive Shoots of Grassland Species from North-Eastern China

The differences in net photosynthetic rate (P _N), transpiration rate (E), and water use efficiency (WUE) between the vegetative and reproductive shoots of three native grass species from the grassland of northeastern China [grey-green and yellow green populations of Leymus chinensis (Trin.) Tzvel., Puccinellia tenuiflora (Griseb) Scrib & Merr, Puccinellia chinampoensis Ohwi] were compared. The two type shoots experienced similar habitats, but differed in leaf life-span and leaf area. The leaf P _N and WUE for the vegetative shoots were significantly higher than those for the reproductive shoots in the grasses, while their E were remarked lower in the dry season. Relative lower leaf P _N and WUE for the reproductive shoots of grassland grasses may explain the facts of lower seed production and the subordinate role of seed in the grassland renewal in north-eastern China.     

Effects of defoliation and girdling on fruit production in Ilex aquifolium

1. The effects of defoliation and girdling on fruit production in European holly Ilex aquifolium were studied in a northern Spanish population. Three defoliation treatments (control, 50% and 100% leaf removal) were applied at branch level in 10 trees. Six branches were used per treatment in each tree and half of these branches were girdled (a ring of bark and cambium was removed from the branch base).
2. Leaf removal from ungirdled branches had little effect on fruit set, fruit production and reproductive allocation at branch level. However, these variables decreased as the extent of defoliation increased on girdled branches.
3. Fruit production in undefoliated branches did not differ between girdled and ungirdled branches, showing that girdled branches were autonomous for fruit production when undamaged. Mean fruit mass of girdled-100% defoliated branches averaged 8% of fruit mass produced by control branches.
4. Fruit production of ungirdled-100% defoliated branches did not differ from control branches, showing the importance of resource importation from other branches. Hence the branches may be regarded as only partially autonomous for fruit production.
5. Fruit production of girdled-50% defoliated branches was 42% of fruit production of control branches, while ungirdled-50% defoliated branches amounted to 70% of controls. These results also demonstrated resource importation but did not support the existence of photosynthetic compensatory mechanisms.
6. The ability of resource importation developed by fruiting branches might be a mechanism to reduce the effect of reproductive costs in this species.     

How does Dryobalanops aromatica supply carbohydrate resources for reproduction in a masting year?

The effect on reproduction of the dynamics of resource allocation was studied in an emergent and masting tree species, Dryobalanops aromatica (Dipterocarpaceae), in a lowland dipterocarp forest in Sarawak, Malaysia. Girdling of the reproductive shoots (5 mm diameter) caused an increase in abortion during the flowering period, but did not affect the fruit set at the middle or final stages of seed maturation. In contrast, 50% defoliation significantly affected fruit setting, but had little effect on flowering. The total leaf area of reproductive shoots was significantly correlated with final fruit set and total fruit mass. Control of the carbohydrate supply to reproductive shoots by girdling and defoliation made no difference to fruit size, but the fruit number was highly sensitive to carbohydrate availability. Total non-structural carbohydrate (TNC) decreased during the flowering period mainly in the branch (P<0.05), but fluctuated little in any organs during fruit maturation. Leaf nitrogen and photosynthetic capacity of the reproductive shoots were not significant variables for reproduction. Our results suggest that D. aromatica uses current photosynthates in the leaves of reproductive shoots as a carbon source during fruit development, but requires stored assimilates in the branch for flowering. However, since TNC was still present in all organs even after flowering, our study also suggests that storage of carbohydrate resources might not be the decisive factor in the occurrence or frequency of flowering in this species.     

Net CO(2) Assimilation and Carbohydrate Partitioning of Grapevine Leaves in Response to Trunk Girdling and Gibberellic Acid Application

Leaf net CO₂ assimilation rate (A), stomatal conductance (g_s), carboxylation efficiency, and foliar nonstructural carbohydrates were measured on mature, field-grown Vitis vinifera L. (cv Thompson Seedless) vines that had been trunk girdled, sprayed with gibberellic acid, or both, shortly after anthesis. Girdling reduced A, g_s, and carboxylation efficiency when measured 2 weeks after imposition of the treatments. Diurnal measurements indicated that A of girdled vines was less than that of control vines between 1000 and 1800 hours. Gibberellic acid mitigated the depressing effect of girdling on g_s during the same diurnal measurements. The concentrations of foliar carbohydrates were greatest for the girdled vines, followed by the combination treatment and were lowest for the control vines. Foliar carbohydrates were greater for girdled vines 4 weeks after the treatments were imposed, however, by this time there was no significant difference in A between the control and girdled vines. Two and 4 weeks after the experiment was initiated root carbohydrate concentrations were less for the girdled vines when compared to the control vines. The data indicate that the reduction in A of girdled grapevines is not associated with the accumulation of leaf nonstructural carbohydrates following the girdling treatment.     

Rearrangement of leaf traits with changing source-sink relationship in blueberry (Vaccinium corymbosum L.) leaves

The source-sink relationship is one of major determinants of plant performance. The influence of reproductive sink demand on light-saturated photosynthesis (P_max), dark respiration (R_D), stomatal conductance (g_s), intrinsic water-use efficiency (WUE_i), contents of soluble sugar (SSC), nitrogen, carbon, and photosynthetic pigments was examined in blueberry (Vaccinium corymbosum L. cv. ‘Brigitta’) during the final stage of rapid fruit growth. Measurements were performed three times per day on developed, sun-exposed leaves of girdled shoots with 0.1, 1, and 10 fruit per leaf (0.1F:L, 1F:L, and 10F:L, respectively) and nongirdled shoots bearing one fruit per leaf (NG). Girdling and lower fruit amount induced lower P_max, g_s, N, and total chlorophyll (Chl) and higher WUE_i, SSC, R_D, Chl a/b ratio and carotenoids-to-chlorophylls ratio (Car/Chl) for the 1F:L and 0.1F:L treatments. The impact of girdling was counterbalanced by 10F:L, with NG and 10F:L having similar values. Variables other than P_max, R_D, g_s, WUE_i, and SSC were unaffected throughout the course of the day. P_max and g_s decreased during the course of the day, but g_s decreased more than P_max in the afternoon, while WUE_i was increasing in almost all treatments. SSC increased from the morning until afternoon, whereas R_D peaked at noon regardless of the treatment. Generally, P_max was closely and negatively correlated to SSC, indicating that sugar-sensing mechanisms played an important role in regulation of blueberry leaf photosynthesis. With respect to treatments, P_max and N content were positively related, while R_D was not associated to substrate availability. The enhanced Car/Chl ratio showed a higher photoprotection under the lower sink demand. Changes in the source-sink relationship in ‘Brigitta’ blueberry led to a rearrangement of physiological and structural leaf traits which allowed adjusting the daily balance between carbon assimilation and absorbed light energy.     

Photosynthesis and chlorophyll fluorescence response to low sink demand of tubers and roots in <Emphasis Type="Italic">Dahlia pinnata</Emphasis> source leaves

Photosynthetic rate (P_N) and chlorophyll (Chl) fluorescence induction of source leaves in response to a low sink demand created by girdling the branch (GB) between the root-tuber-system and the leaves were studied in Dahlia pinnata L. cv. Rigolet during the stage of rapid tuber growth in the greenhouse. GB resulted in significantly lower values of P_N, stomatal conductance (g_s), and transpiration rate (E), but in higher leaf temperature (T_l) compared with those of controls. With exception of maximum quantum yield of photosystem 2 (PS 2) photochemistry (F_v/F_m) and maximum ratio of quantum yields of photochemical and concurrent non-photochemical processes in PS 2 (F_v/F₀), no significant differences were observed in Chl fluorescence parameters between girdled and control leaves on days 1 and 2 after GB, indicating no apparent damage in the photosynthetic apparatus. However, longer girdling duration resulted in higher non-photochemical Chl fluorescence quenching (NPQ), but lower F_v/F₀, actual efficiency of energy conversion in PS 2 under steady-state conditions (Φ_PS2), and photochemical quenching coefficient (qP) in comparison with controls from 10:00 to 16:00 or 15:00 on days 4 and 5, respectively, indicating reversible injury in the photosynthetic apparatus.     

Physiological divergences between two rhizomatous grasses from a desertification steppe,North China

The inter-and intra-specific physiological differences, e.g. rates of net photosynthesis (P _N) and transpiration (E), stomatal conductance (g _s), and water use efficiency (WUE), were compared between two grasses, Calamagrostis epigeios (L.) Roth. and Psammochloa villosa (Trin.) Bor., and between their leaf types in a desertification steppe in North China. The two species had a similar habitat, but differed in leaf area and rhizome depth. Leaf P _N, E, and g _s for P. villosa were significantly greater than those for C. epigeios in the growing season, but WUE for the former species was only 50 and 80 % of that for the latter one in dry and rainy seasons, respectively. In general, leaf P _N, E, g _s, and WUE for both vegetative and reproductive shoots of the two species exhibited little variations between leaf types or with leaf age, even though there were some remarkable differences between dry and rainy seasons. The mean leaf P _N and E in reproductive shoots of P. villosa were significantly lower than those in its vegetative shoots in rainy season, while these differences were much smaller for those of C. epigeios. P. villosa with deeper rhizome roots has relative higher leaf P _N, E, and g _s, but a smaller WUE in the arid desertification steppe region.     

Stem girdling influences concentrations of endogenous cytokinins and abscisic acid in relation to leaf senescence in cotton

Many studies have shown that root–shoot imbalance influences vegetative growth and development of cotton (Gossypium hirsutum L.), but few have examined changes in leaf senescence and endogenous hormones due to stem girdling. The objective of this study was to determine the correlation between some endogenous phytohormones, particularly cytokinins and abscisic acid (ABA), and leaf senescence following stem girdling. Field-grown cotton plants were girdled on the main stem 5 days after squaring (DAS), while the non-girdled plants served as control. Plant biomass, seed cotton yield, main-stem leaf photosynthetic (Pn) rate, chlorophyll (Chl) and malondialdehyde (MDA) concentrations, as well as levels of cytokinins and ABA in main-stem leaves and xylem sap were determined after girdling or at harvest. Main-stem girdling decreased the dry root weight and root/shoot ratio from 5 to 70 days after girdling (DAG) and reduced seed cotton yield at harvest. Main-stem leaf Pn and Chl concentration in girdled plants were significantly lower than in control plants. Much higher levels of MDA were observed in main-stem leaves from 5 to 70 DAG, suggesting that stem girdling accelerated leaf senescence. Girdled plants contained less trans-zeatin and its riboside (t-Z + t-ZR), dihydrozeatin and its riboside (DHZ + DHZR), and isopentenyladenine and its riboside (iP + iPA), but more ABA than control plants in both main-stem leaves and xylem sap. These results suggested that main-stem girdling accelerated leaf senescence due to reduced levels of cytokinin and/or increased ABA. Cytokinin and ABA are involved in leaf senescence following main-stem girdling.     

Variations in photosynthetic rate and associated parameters with age of oil palm leaves under irrigation

Net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) in an adult oil palm (Elaeis guineensis) canopy were highest in the 9^th leaf and progressively declined with leaf age. Larger leaf area (LA) and leaf dry mass (LDM) were recorded in middle leaves. P _N showed a significant positive correlation with g _s and a negative relationship with leaf mass per area (ALM). The oil palm leaf remains photosynthetically active for a longer time in the canopy which contributes significantly to larger dry matter production in general and greater fresh fruit bunch yields in particular.     

Girdling induces oxidative damage and triggers enzymatic and non-enzymatic antioxidative defences in Citrus leaves

The effects of girdling on oxidative damage, antioxidant enzyme activity, antioxidant metabolites and proline (Pro) were studied in leaves arising from different shoot types of potted 2-year-old ‘Loretina’ mandarin (Citrus reticulata Blanco) trees during the spring flush period. Girdling increased malonyldialdehyde (MDA) and basal chlorophyll (Chl) a fluorescence (F_o) in young leaves 30 days after girdling but not in the mature leaves (ML) suggesting a disruption of photosynthetic apparatus and oxidative damage in young leaves. This phenomenon was accompanied by increasing levels of Pro. Paralleling these changes, an increase of all antioxidant enzyme activities occurred in leaves from vegetative (VG) and multiflowered leafy shoots (MLY) of girdled trees. Similarly, in ML of girdled trees, ascorbate peroxidase (APX), catalase (CAT) and glutathione reductase (GR) activity also increased. However, dehydroascorbate reductase (DHAR) activity decreased and superoxide dismutase (SOD) activity remained unchanged. Total leaf carbohydrate content and starch also increased as a result of girdling in all shoot types. Whilst soluble sugars increased markedly in young leaves, they increased only slightly in ML. In conclusion, this study provides evidence that girdling gives rise to oxidative damage in Citrus during carbohydrate accumulation, triggering enzymatic and non-enzymatic defence mechanisms.     

Photosynthesis,Transpiration, and Water Use Efficiency of Two Puccinellia Species on the Songnen Grassland,Northeastern China

The intra- and inter-specific variations in net photosynthetic (P _N) and transpiration (E) rates and water use efficiency (WUE) of Puccinellia tenuiflora and Puccinellia chinampoensis leaves were compared. The two species experienced a similar habitat, but differed in leaf area, leaf colour, and nitrogen contents. Leaf P _N and E for both reproductive and vegetative shoots of the two species declined with leaf age. P _N for reproductive shoots was less than for vegetative shoots, but their E was greater than that of vegetative shoots in the dry season. The average P _N and E for reproductive shoots of P. tenuiflora were lower than those of P. chinampoensis, but higher for vegetative shoots.     

Girdling induces oxidative damage and triggers enzymatic and non-enzymatic antioxidative defences in Citrus leaves

The effects of girdling on oxidative damage, antioxidant enzyme activity, antioxidant metabolites and proline (Pro) were studied in leaves arising from different shoot types of potted 2-year-old ‘Loretina’ mandarin (Citrus reticulata Blanco) trees during the spring flush period. Girdling increased malonyldialdehyde (MDA) and basal chlorophyll (Chl) a fluorescence (F_o) in young leaves 30 days after girdling but not in the mature leaves (ML) suggesting a disruption of photosynthetic apparatus and oxidative damage in young leaves. This phenomenon was accompanied by increasing levels of Pro. Paralleling these changes, an increase of all antioxidant enzyme activities occurred in leaves from vegetative (VG) and multiflowered leafy shoots (MLY) of girdled trees. Similarly, in ML of girdled trees, ascorbate peroxidase (APX), catalase (CAT) and glutathione reductase (GR) activity also increased. However, dehydroascorbate reductase (DHAR) activity decreased and superoxide dismutase (SOD) activity remained unchanged. Total leaf carbohydrate content and starch also increased as a result of girdling in all shoot types. Whilst soluble sugars increased markedly in young leaves, they increased only slightly in ML. In conclusion, this study provides evidence that girdling gives rise to oxidative damage in Citrus during carbohydrate accumulation, triggering enzymatic and non-enzymatic defence mechanisms.