Effects of Water Stress on Photosynthesis and Carbon Partitioning in Soybean (Glycine max [L.] Merr.) Plants Grown in the Field at Different CO(2) Levels

The effects of water stress and CO₂ enrichment on photosynthesis, assimilate export, and sucrose-P synthase activity were examined in field grown soybean plants. In general, leaves of plants grown in CO₂-enriched atmospheres (300 microliters per liter above unenriched control, which was 349 ± 12 microliters per liter between 0500 and 1900 hours EST over the entire season) had higher carbon exchange rates (CER) compared to plants grown at ambient CO₂, but similar rates of export and similar activities of sucrose-P synthase. On most sample dates, essentially all of the extra carbon fixed as a result of CO₂ enrichment was partitioned into starch. CO₂-enriched plants had lower transpiration rates and therefore had a higher water use efficiency (milligrams CO₂ fixed per gram H₂O transpired) per unit leaf area compared to nonenriched plants. Water stress reduced CER in nonenriched plants to a greater extent than in CO₂-enriched plants. As CER declined, stomatal resistance increased, but this was not the primary cause of the decrease in assimilation because internal CO₂ concentration remained relatively constant. Export of assimilates was less affected by water stress than was CER. When CERs were low as a result of the imposed stress, export was supported by mobilization of reserves (mainly starch). Export rate and leaf sucrose concentration were related in a curvilinear manner. When sucrose concentration was above about 12 milligrams per square decimeter, obtained with nonstressed plants at high CO₂, there was no significant increase in export rate. Assimilate export rate was also correlated positively with SPS activity and the quantitative relationship varied with CER. Thus, export rate was a function of both CER and carbon partitioning.     

Diurnal changes in maize leaf photosynthesis : I. Carbon exchange rate, assimilate export rate, and enzyme activities

Diurnal changes in photosynthetic parameters and enzyme activities were characterized in greenhouse grown maize plants (Zea mays L. cv Pioneer 3184). Rates of net photosynthesis and assimilate export were highest at midday, coincident with maximum irradiance. During the day, assimilate export accounted for about 80% of net carbon fixation, and the maximum export rate (35 milligrams CH₂O per square decimeter per hour) was substantially higher than the relatively constant rate maintained through the night (5 milligrams CH₂O per square decimeter per hour). Activities of sucrose phosphate synthase and NADP-malate dehydrogenase showed pronounced diurnal fluctuations; maximum enzyme activities were generally coincident with highest light intensity. Reciprocal light/dark transfers of plants throughout the diurnal cycle revealed that both enzymes were deactivated by 30 minutes of darkness during the day, and they could both be substantially activated by 30 minutes of illumination at night. During 24 hours of extended darkness, sucrose phosphate synthase activity declined progressively to an almost undetectable level, but was activated after 1.5 hours of illumination. Thus, the diurnal fluctuation in maize sucrose phosphate synthase can be explained by some form of light modulation of enzyme activity and is not due to an endogenous rhythm in activity. No diurnal fluctuations were observed in the activities of NADP-malic enzyme or fructose 6-phosphate-2-kinase. Phosphoenolpyruvate carboxylase was activated by light to some extent (about 50%) when activity was measured under suboptimal conditions in vitro. The results suggested that the rates of sucrose formation and assimilate export were closely aligned with the rate of carbon fixation and the activation state of sucrose phosphate synthase.     

Carbon assimilation,partitioning and export in mature cladophylls of two asparagus (Asparagus officinalis) cultivars with contrasting yield

To assess the physiological aspects underpinning cultivar difference in asparagus ( Asparagus officinalis L.) yield, diel carbon exchange, carbon partitioning and export and sucrose phosphate synthase (SPS) activity were examined in mature cladophyll tissue of two asparagus cultivars (ASP-69 and ASP-03) under field conditions. Both cultivars exhibited similar diel patterns in carbon exchange rate (CER) and carbohydrate partitioning. Rates of carbon export estimated from CER and dry mass changes were the highest at midday and coincided with maximum assimilation rate. Carbon export accounted for about 74% of carbon fixation during the photoperiod. No diel fluctuations were observed in SPS activity in either cultivar. A positive correlation between day CER and carbon export rate (r² = 0.87) was found and this relationship did not differ between the two cultivars studied. The greater carbon export rate measured in the high-yielding cultivar (ASP-69) was associated with significantly higher CER in comparison to the low-yielding cultivar (ASP-03). However, a correlation between sucrose concentration and carbon export rate did not exist. Biochemical evidence indicated that the greater CER in ASP-69 was associated with a significantly greater SPS activity ( P   0.05). Phloem ¹⁴C exudate analysis revealed that ¹⁴C flux out of cladophyll tissue in ASP-69 was significantly greater than in ASP-03. These results indicate a feed-forward effect of rate of photosynthesis on assimilate export in the two cultivars studied.     

Regulation of photosynthetic carbon metabolism in cucumber by light intensity and photosynthetic period

The effects of photosynthetic periods and light intensity on cucumber (Cucumis sativus L.) carbon exchange rates and photoassimilate partitioning were determined in relation to the activities of galactinol synthase and sucrose-phosphate synthase. Carbon assimilation and partitioning appeared to be controlled by different mechanisms. Carbon exchange rates were influenced by total photon flux density, but were nearly constant over the entire photoperiod for given photoperiod lengths. Length of the photosynthetic periods did influence photoassimilate partitioning. Assimilate export rate was decreased by more than 60% during the latter part of the short photoperiod treatment. This decrease in export rate was associated with a sharp increase in leaf starch acccumulation rate. Results were consistent with the hypothesis that starch accumulation occurs at the expense of export under short photoperiods. Galactinol synthase activities did not appear to influence the partitioning of photoassimilates between starch and transport carbohydrates. Sucrose phosphate synthase activities correlated highly with sugar formation rates (sucrose, raffinose, stachyose + assimilate export rate, r = 0.93, α = 0.007). Cucumber leaf sucrose phosphate synthase fluctuated diurnally in a similar pattern to that observed in vegetative soybean plants.     

Diurnal fluctuations in cotton leaf carbon export, carbohydrate content, and sucrose synthesizing enzymes

In fully expanded leaves of greenhouse-grown cotton (Gossypium hirsutum L., cv Coker 100) plants, carbon export, starch accumulation rate, and carbon exchange rate exhibited different behavior during the light period. Starch accumulation rates were relatively constant during the light period, whereas carbon export rate was greater in the afternoon than in the morning even though the carbon exchange rate peaked about noon. Sucrose levels increased throughout the light period and dropped sharply with the onset of darkness; hexose levels were relatively constant except for a slight peak in the early morning. Sucrose synthase, usually thought to be a degradative enzyme, was found in unusually high activities in cotton leaf. Both sucrose synthase and sucrose phosphate synthetase activities were found to fluctuate diurnally in cotton leaves but with different rhythms. Diurnal fluctuations in the rate of sucrose export were generally aligned with sucrose phosphate synthase activity during the light period but not with sucrose synthase activity; neither enzyme activity correlated with carbon export during the dark. Cotton leaf sucrose phosphate synthase activity was sufficient to account for the observed carbon export rates; there is no need to invoke sucrose synthase as a synthetic enzyme in mature cotton leaves. During the dark a significant correlation was found between starch degradation rate and leaf carbon export. These results indicate that carbon partitioning in cotton leaf is somewhat independent of the carbon exchange rate and that leaf carbon export rate may be linked to sucrose formation and content during the light period and to starch breakdown in the dark.     

Characterization of diurnal changes in activities of enzymes involved in sucrose biosynthesis

Experiments were conducted with vegetative soybean plants (Glycine max [L.] Merr., `Ransom') to determine whether the activities in leaf extracts of key enzymes in sucrose metabolism changed during the daily light/dark cycle. The activity of sucrose-phosphate synthase (SPS) exhibited a distinct diurnal rhythm, whereas the activities of UDP-glucose pyrophosphorylase, cytoplasmic fructose-1,6-bisphosphatase, and sucrose synthase did not. The changes in extractable SPS activity were not related directly to photosynthetic rates or light/dark changes. Hence, it was postulated that the oscillations were under the control of an endogenous clock. During the light period, the activity of SPS was similar to the estimated rate of sucrose formation. In the dark, however, SPS activity declined sharply and then increased even though degradation of starch was linear. The activity of SPS always exceeded the estimated maximum rate of sucrose formation in the dark. Transfer of plants into light during the normal dark period (when SPS activity was low) resulted in increased partitioning of photosynthate into starch compared to partitioning observed during the normal light period. These results were consistent with the hypothesis that SPS activity in situ was a factor regulating the rate of sucrose synthesis and partitioning of fixed carbon between starch and sucrose in the light.     

Effect of Photoperiod on Photosynthate Partitioning and Diurnal Rhythms in Sucrose Phosphate Synthase Activity in Leaves of Soybean (Glycine max L. [Merr.]) and Tobacco (Nicotiana tabacum L.)

Studies were conducted to identify the existence of diurnal rhythms in sucrose phosphate synthase (SPS) activity in leaves of three soybean (Glycine max L. [Merr.]) and two tobacco (Nicotiana tabacum L.) cultivars and the effect of photoperiod (15 versus 7 hours) on carbohydrate partitioning and the rhythm in enzyme activity. Acclimation of all the genotypes tested to a short day (7 hours) photoperiod resulted in increased rates of starch accumulation, whereas rates of translocation, foliar sucrose concentrations, and activities of SPS were decreased relative to plants acclimated to long days (15 hours). Under the long day photoperiod, two of the three soybean cultivars (`Ransom' and `Jupiter') and one of the two tobacco cultivars (`22NF') studied exhibited a significant diurnal rhythm in SPS activity. With the soybean cultivars, acclimation to short days reduced the activity of SPS (leaf fresh weight basis) and tended to dampen the amplitude of the rhythm. With the tobacco cultivars, photoperiod affected the shape of the SPS-activity rhythm. The mean values for SPS activity (calculated from observations made during the light period) were correlated positively with translocation rates and were correlated negatively with starch accumulation rates. Overall, the results support the postulate that SPS activity is closely associated with starch/sucrose levels in leaves, and that acclimation to changes in photoperiod may be associated with changes in the activity of SPS.     

Trends in carbohydrate depletion, respiratory carbon loss, and assimilate export from soybean leaves at night

To evaluate assimilate export from soybean (Glycine max [L.] Merrill) leaves at night, rates of respiratory CO₂ loss, specific leaf weight loss, starch mobilization, and changes in sucrose concentration were measured during a 10-hour dark period in leaves of pod-bearing `Amsoy 71' and `Wells II' plants in a controlled environment. Lateral leaflets were removed at various times between 2200 hours (beginning dark period) and 0800 hours (ending dark period) for dry weight determination and carbohydrate analyses. Respiratory CO₂ loss was measured throughout the 10-hour dark period. Rate of export was estimated from the rate of loss in specific leaf weight and rate of CO₂ efflux. Rate of assimilate export was not constant. Rate of export was relatively low during the beginning of the dark period, peaked during the middle of the dark period, and then decreased to near zero by the end of darkness. Rate of assimilate export was associated with rate of starch mobilization and amount of starch reserves available for export. Leaves of Amsoy 71 had a higher maximum export rate in conjunction with a greater total change in starch concentration than did leaves of Wells II. Sucrose concentration rapidly declined during the first hour of darkness and then remained constant throughout the rest of the night in leaves of both cultivars. Rate of assimilate export was not associated with leaf sucrose concentration.     

Diurnal Changes in Maize Leaf Photosynthesis : II. Levels of Metabolic Intermediates of Sucrose Synthesis and the Regulatory Metabolite Fructose 2,6-Bisphosphate

Diurnal changes in the regulatory metabolite, fructose-2,6-bisphosphate (F26BP), and key metabolic intermediates of sucrose biosynthesis were studied in maize (Zea mays L. cv Pioneer 3184) during a day-night cycle. Whole leaf concentrations of dihydroxyacetonephosphate (DHAP) and fructose 1,6-bisphosphate changed markedly during the photoperiod. DHAP concentration was correlated positively with the rate of sucrose formation in vivo (assimilate export plus sucrose accumulation) and extractable activity of sucrose phosphate synthase (SPS). The changes closely followed net photosynthetic rate, which tracked irradiance. The other metabolic intermediates measured (glucose 6-phosphate, fructose 6-phosphate, and UDP-glucose) were either relatively constant over the 24 hour period or changed in a different pattern. Diurnal changes in leaf F26BP concentrations were pronounced, and fundamentally different than the pattern reported with other species. F26BP concentration decreased at the beginning of the day and remained low and constant; a 3- to 4-fold increase occurred with darkness, and slowly declined thereafter. In general, leaf F26BP concentration was negatively correlated with net photosynthetic rate, and also leaf DHAP concentration. Consequently, co-ordination of the regulation of cytosolic fructose 1,6-bisphosphatase and SPS was apparent. The results support the postulate that in maize leaves the activation state of SPS may be dependent on availability of DHAP and possibly other metabolites.     

Effects of Low Temperature on Photosynthesis in Flag Leaves of Rice (Oryza sativa L.)

After low temperature (16℃) treatment, the net photosynthetic rate, chlorophyll content and FBPase activity tended to decrease and ratio of chlorophyll a/b, stomatal resistance, sucrose and starch content tended to increase in rice flag leaves at heading stage. However, the Pi content and water potential did not show pronounced change. The flag leaf chloroplasts contained large starch grains after three days at low temperature. The recovery of net photosynthetic rate in the flag leaves which had recieved stress at 16℃ for three days and transferred to normal nature condition was gradually increased. It almost came up to the value of control after five days. However, the decrease in stomatal resistance was rapid. It was almost lowered to the value of control after one day. Under normal nature comdition for five days, the chlorophyll content and FBPase activity reached near to the value of control. Meanwhile, the ratio of chlorophyll a/b, the sucrose and starch content almost declined to the value of control. The large starch grain in flag leaf chloroplast also disappeared. According to the experimental results, we discussed the relationship between the above-mentioned various changes and net photosynthetic rate We suggested that the effect of low temperature on photosynthesis might be one of factors in yield-reduction of rice.     

Regulation of photosynthesis by end-product accumulation in leaves of plants storing starch, sucrose, and hexose sugars

In the present study, leaves of different plant species were girdled by the hot wax collar method to prevent export of assimilates. Photosynthetic activity of girdled and control leaves was evaluated 3 to 7 days later by two methods: (a) carbon exchange rate (CER) of attached leaves was determined under ambient CO₂ concentrations using a closed gas system, and (b) maximum photosynthetic capacity (A_max) was determined under 3% CO₂ with a leaf disc O₂ electrode. Starch, hexoses, and sucrose were determined enzymically. Typical starch storers like soybean (Glycine max L.) (up to 87.5 milligrams of starch per square decimeter in girdled leaves), cotton (Gossypium hirsutum L.), and cucumber (Cucumis sativus L.) responded to 7 days of girdling by increased (80-100%) stomatal resistance (r_s) and decreased A_max (>50%). On the other hand, spinach (Spinacia oleracea L.), a typical sucrose storer (up to 160 milligrams of sucrose per square decimeter in girdled leaves), showed only a slight reduction in CER and almost no change in A_max. Intermediate plants like tomato (Lycopersicon esculentum Mill.), sunflower (Helianthus annuus L.), broad bean (Vicia faba L.), bean (Phaseolus vulgaris L.), and pea (Pisum sativum L.), which upon girdling store both starch and sucrose, responded to the girdle by a considerable reduction in CER but only moderate inhibition of A_max, indicating that the observed reduction in CER was primarily a stomatal response. Both the wild-type tobacco (Nicotiana sylvestris) (which upon girdling stored starch and hexoses) and the starchless mutant (which stored only hexoses, up to 90 milligrams per square decimeter) showed 90 to 100% inhibition of CER and approximately 50% inhibition of A_max. In general, excised leaves (6 days) behaved like girdled leaves of the respective species, showing 50% reduction of A_max in wild-type and starchless N. sylvestris but only slight decline of A_max in spinach. The results of the present study demonstrate the possibility of the occurrence of end-product inhibition of photosynthesis in a large number of crop plants. The long-term inhibition of photosynthesis in girdled leaves is not confined to stomatal responses since the A_max declined up to 50%. The inhibition of A_max by girdling was strongest in starch storers, but starch itself cannot be directly responsible, because the starchless mutant of N. sylvestris was also strongly inhibited. Similarly, the inhibition cannot be attributed to hexose sugars either, because soybean, cotton, and cucumber are among the plants most strongly inhibited although they do not maintain a large hexose pool. Spinach, a sucrose storer, showed the least inhibition in both girdled and excised leaf systems, which indicates that sucrose is probably not directly responsible for the end-product inhibition of photosynthesis. The occurrence of strong end-product inhibition appears to be correlated with high acid-invertase activity in fully expanded leaves. The inhibition may be related to the nature of soluble sugar metabolism in the extrachloroplastic compartment and may be caused by a metabolite that has different rates of accumulation and turnover in sucrose storers and other plants.     

Biochemical basis for effects of k-deficiency on assimilate export rate and accumulation of soluble sugars in soybean leaves

The effects of K-deficiency on carbon exchange rates (CER), photosynthate partitioning, export rate, and activities of key enzymes involved in sucrose metabolism were studied in soybean (Glycine max [L.] Merr.) leaves. The different parameters were monitored in mature leaves that had expanded prior to, or during, imposition of a complete K-deficiency (plants received K-free nutrition solution). In general, recently expanded leaves had the highest concentration of K, and imposition of K-stress at any stage of leaf expansion resulted in decreased K concentrations relative to control plants (10 millimolar K). A reduction in CER, relative to control plants, was only observed in leaves that expanded during the K-stress. Stomatal conductance also declined, but this was not the primary cause of the decrease in carbon fixation because internal CO₂ concentration was unaffected by K-stress. Assimilate export rate from K-deficient leaves was reduced but relative export, calculated as a percentage of CER, was similar to control leaves. Over all the data, export rate was correlated positively with both CER and activity of sucrose phosphate synthase in leaf extracts. K-deficient leaves had higher concentrations of sucrose and hexose sugars. Accumulation of hexose sugars was associated with increased activities of acid invertase. Neutral invertase activity was low and unaffected by K-nutrition. It is concluded that decreased rates of assimilate export are associated with decreased activities of sucrose phosphate synthase, a key enzyme involved in sucrose formation, and that accumulation of hexose sugars may occur because of increased hydrolysis of sucrose in K-deficient leaves.     

去果河套蜜瓜源叶碳水化合物及其相关酶昼夜变化特征

以河套蜜瓜为试材,在果实迅速膨大期通过去果处理改变库源关系,研究源叶净光合速率,蔗糖、还原糖和淀粉含量及其代谢相关酶活性的昼夜变化规律。结果表明:(1)源叶的净光合速率为单峰曲线,无明显的"光合午休"现象,去果处理对其无影响。(2)源叶中蔗糖和还原糖含量的昼夜变化为单峰曲线,蔗糖磷酸合成酶和蔗糖合成酶合成方向活性的昼夜变化为双峰曲线,蔗糖合成酶分解方向、酸性转化酶和中性转化酶活性的昼夜变化无明显规律,改变库源关系对这些指标均无显著影响;蔗糖含量升高受蔗糖磷酸合成酶和蔗糖合成酶合成方向正调控,而蔗糖含量降低则受多种酶的共同调节。(3)源叶中淀粉含量和腺苷二磷酸葡萄糖焦磷酸化酶活性的昼夜变化为单峰曲线,去果处理可以显著提高淀粉含量和腺苷二磷酸葡萄糖焦磷酸化酶活性,淀粉含量升高受腺苷二磷酸葡萄糖焦磷酸化酶正调控。     

Daytime and nighttime carbon balance and assimilate export in soybean leaves at different photon flux densities

To evaluate daytime and nighttime carbon balance and assimilate export in soybean (Glycine max [L.] Merrill) leaves at different photon flux densities, rates of CO₂ exchange, specific leaf weights, and concentrations of sucrose and starch were measured at intervals in leaves of pod-bearing `Amsoy 71' and `Wells II' plants grown in a controlled environment room. Assimilate export was estimated from CO₂ exchange and change in specific leaf weight. Total diurnal assimilate export was similar for both cultivars. Large cultivar differences existed, however, in the partitioning of carbon into starch reserves and the relative amounts of assimilate exported during the day and the night. Total amounts of both daytime and nighttime export increased with increasing photon flux density, as did sucrose and starch concentrations, specific leaf weight, and rate of respiratory carbohydrate loss at night. Cultivar differences in nighttime rate of export were more closely related to the differences in amount of assimilate available at the end of the day than to differences in daytime rate of net CO₂ assimilation. Daytime rates of export, however, were closely related to daytime rates of net CO₂ assimilation within each cultivar. The total amount of starch depleted during the 10-hour night increased as starch concentration at the beginning of the night increased.     

The effect of water stress on photosynthetic carbon metabolism in four species grown under field conditions

Abstract. The effect of gradually-developing water-stress has been studied in Lupinus albus L., Helianthus annuus L., Vitis vinifera cv. Rosaki and Eucalyptus globulus Labill. Water was withheld and diurnal rhythms were investigated 4–8d later, when the predawn water deficit was more negative than in watered plants, and the stomata closed almost completely early during the photoperiod. The contribution of ‘stomatal’ and ‘non-stomatal’ components to the decrease of photosynthetic rate was investigated by (1) comparing the changes of the rate of photosynthesis in air with the changes of stomatal conductance and (2) measuring photosynthetic capacity in saturating irradiance and 15% CO₂. Three species (lupin, eucalyptus and sunflower) showed larger changes of stomatal conductance than photosynthesis in air, and showed little or no decrease of photosynthetic capacity in saturating CO₂. Photosynthesis in air also recovered fully overnight after watering the plants in the evening. In grapevines, stomatal conductance and photosynthesis in air changed in parallel, there was a marked decrease of photosynthetic capacity, and photosynthesis and stomatal conductance did not recover overnight after watering water-stressed plants. Relative water content remained above 90% in grapevine. We conclude that non-stomatal components do not play a significant role in lupins, sunflower or eucalyptus, but could in grapevine. The effect of water-stress on partitioning of photosynthate was investigated by measuring the amounts of sucrose and starch in leaves during a diurnal rhythm, and by measuring the partitioning of ¹⁴C-carbon dioxide between sucrose and starch. In all four species, starch was depleted in water-stressed leaves but sucrose was maintained at amounts similar to, or higher than, those in watered plants. Partitioning into sucrose was increased in lupins and eucalyptus, and remained unchanged in grapevine and sunflower. It is concluded that water-stressed leaves in all four species maintain high levels of soluble sugars in their leaves, despite having lower rates of field photosynthesis, decreased rates of export, and low amounts of starch in their leaves.     

Coarse control of sucrose-phosphate synthase in leaves: Alterations of the kinetic properties in response to the rate of photosynthesis and the accumulation of sucrose

It has been investigated whether diurnal rhythms of sucrose-phosphate synthase (SPS) are involved in controlling the rate of photosynthetic sucrose synthesis. Extracts were prepared from spinach (Spinacia oleracea L.) and barley (Hordeum vulgare L.) leaves and assayed for enzyme activity. The activity of SPS increased in parallel with a rising rate of photosynthesis, and was increased by feeding mannose and decreased by supplying inorganic phosphate. In leaf material where sucrose had accumulated during the photoperiod or when sucrose was supplied exogenously, SPS activity decreased. During a diurnal rhythm, SPS activity increased after illumination, declined gradually during the light period, decreased further after darkening and then recovered gradually during the night. These changes did not involve an alteration of the maximal activity, but were caused by changes in the kinetic properties, revealed as a change in sensitivity to inhibition by inorganic phosphate. In experiments which modelled the response of SPS to changing metabolite concentrations, it was shown that these alterations of kinetic properties would strongly modify the activity of SPS in vivo. It is proposed that SPS can exist in kinetically distinct forms in vivo, and that the distribution between these forms can be rapidly altered. As the rate of photosynthesis increases there is an activation of SPS, which may be directly or indirectly linked to changes in the availability of P_i. This activation can be modified by factors related to the accumulation of sucrose. Under normal conditions there is a balance between these factors, and the leaf contains a mixture of the different forms of SPS.Abbreviations Chl chlorophyll - Frul,6bisP fructose-1,6-bisphosphate - Fru2,6bisP fructose-2,6-bisphosphate - Fru6P fructose-6-phosphate - Fru1,6bisPase fructose-1,6-bisphosphatase - Fru6P 2kinase fructose-6-phosphate, 2kinase - Fru2,6bisPase fructose-2,6-bisphosphatase - Glc6P glucose-6-phosphate - P_j inorganic phosphate - SPS sucrose-phosphate synthase - UDPGLc uridine 5-diphosphate glucose     

Locomotor activity rhythm in two wrasses,Halichoeres tenuispinnis andPteragogus flagellifera,under various light conditions

The locomotor activity rhythms were examined by using an actograph with infra-red photo-electric switches for two species of wrasses, (Halichoeres tenuispinnis andPteragogus flagellifera) under various light conditions. InH. tenuispinnis, the locomotor activity of almost all fish under light-dark cycle regimen (LD12:12; 06:00–18:00 light, 18:00–06:00 dark) commenced somewhat earlier than the beginning of light period and continued till somewhat earlier than the beginning of the dark period. This species clearly showed free-running activity rhythms under both constant illumination (LL) and constant darkness (DD). Therefore,H. tenuispinnis appeared to have a circadian rhythm. The length of the circadian period ranged from 23 hr. 30 min. to 23 hr. 44 min. under LL, and was from 23 hr. 39 min. to 24 hr. 18 min. under DD. On the other hand, the locomotor activity ofP. flagellifera occurred mostly in the light period under LD 12:12. The activity of this species continued through LL, but was greatly suppressed in DD, so that none of the fish had any activity rhythm in both constant conditions. It was known from field observations thatH. tenuispinnis burrowed and lay in sandy bottoms, whileP. flagellifera hid and rested in bases of seagrasses and shallow crevices of rocks during the night. In the present two wrasses, it seemed that the above-mentioned difference of noctural behavior was closely related to the intensity of the endogenous factor in the activity rhythm.     

Studies on Genetic Male-Sterile Soybeans : II. Effect of Nodulation on Photosynthesis and Carbon Partitioning in Leaves

Soybean (Glycine max L. Merr.) germplasm, essentially isogenic except for loci controlling male sterility (ms₁) and nodulation (rj₁), were developed to study the effects of reproductive development and nitrogen source on certain aspects of photosynthesis. Plants were sampled from flowering (77 days after transplanting) until maturity (150 days after transplanting). With all four genotypes, net carbon exchange rates were highest at flowering and declined thereafter. Photosynthetic rates of the sterile genotypes (nodulated and non-nodulated) declined more rapidly than the fertile genotypes, and after 105 days, both sterile genotypes maintained low but relatively constant carbon exchange rates (<3 milligrams CO₂/gram fresh weight per hour). Photosynthetic rates and starch accumulation (difference between afternoon and morning levels) declined with time. The sterile genotypes attained the highest morning starch levels, which reflected reduced starch mobilization. After 92 days, the proportion of photosynthetically fixed carbon that was partitioning into starch (relative leaf starch accumulation) in the sterile genotypes increased dramatically. In contrast, relative leaf starch accumulation in the fertile genotypes remained relatively constant with time. Throughout the test period, all four genotypes maintained leaf sucrose levels between 5 and 15 micromoles glucose equivalents per gram fresh weight.

The activities of sucrose phosphate synthase (SPS) in leaf extracts of the four genotypes declined from 77 to 147 days. Nodulated genotypes tended to maintain higher activities (leaf fresh weight basis) than did the non-nodulated genotypes. In general, relative leaf starch accumulation was correlated negatively with the activity of SPS (normalized with leaf net carbon exchange rate) in leaf extracts for all four genotypes during early reproductive development, and for the fertile genotypes at all sampling dates. In contrast, leaf sucrose content was correlated positively with SPS activity during early reproductive development. These results suggested that a direct relation existed between the activity of SPS and starch/sucrose levels in soybean leaves. However, the interaction between these processes also may be influenced by other factors, particularly when leaf photosynthetic rates and plant demand for assimilates is low, as in the sterile genotypes.

     

Diurnal Changes in Maize Leaf Photosynthesis : III. Leaf Elongation Rate in Relation to Carbohydrates and Activities of Sucrose Metabolizing Enzymes in Elongating Leaf Tissue

Maize (Zea mays L. cv. Pioneer 3184) leaf elongation rate was measured diurnally and was related to diurnal changes in the activities of sucrose metabolizing enzymes and carbohydrate content in the elongating portion of the leaf. The rate of leaf elongation was greatest at midday (1300 hours) and was coincident with the maximum assimilate export rate from the distal portion of the leaf. Leaf elongation during the light period accounted for 70% of the total observed increase in leaf length per 24 hour period. Pronounced diurnal fluctuations were observed in the activities of acid and neutral invertase and sucrose phosphate synthase. Maximum activities of sucrose phosphate synthase and acid invertase were observed at 0900 hours, after which activity declined rapidly. The activity of sucrose phosphate synthase was substantially lower than that observed in maize leaf source tissue. Neutral invertase activity was greatest at midday (1200 hours) and was correlated positively with diurnal changes in leaf elongation rate. There was no significant change in the activity of sucrose synthase over the light/dark cycle. Sucrose accumulation rate increased during a period when leaf elongation rate was maximal and beginning to decline. Maximum sucrose concentration was observed at 1500 hours, when the activities of sucrose metabolizing enzymes were low. At no time was there a significant accumulation of hexose sugars. The rate of starch accumulation increased after the maximum sucrose concentration was observed, continuing until the end of the light period. There was no delay in the onset of starch mobilization at the beginning of the dark period, and essentially all of the starch was depleted by the end of the night. Mobilization of starch in the elongating tissue at night could account for a significant proportion of the calculated increase in the tissue dry weight due to growth. Collectively, the results suggested that leaf growth may be controlled by the activities of certain sucrose metabolizing enzymes and may be coordinated with assimilate export from the distal, source portion of the leaf. Results are discussed with reference to diurnal photoassimilation and export in the distal, source portion of the leaf.     

Decreased expression of two key enzymes in the sucrose biosynthesis pathway, cytosolic fructose-1,6-bisphosphatase and sucrose phosphate synthase, has remarkably different consequences for photosynthetic carbon metabolism in transgenic Arabidopsis thaliana

Photosynthetic carbon metabolism was investigated in antisense Arabidopsis lines with decreased expression of sucrose phosphate synthase (SPS) and cytosolic fructose-1,6-bisphosphatase (cFBPase). In the light, triose phosphates are exported from the chloroplast and converted to sucrose via cFBPase and SPS. At night, starch is degraded to glucose, exported and converted to sucrose via SPS. cFBPase therefore lies upstream and SPS downstream of the point at which the pathways for sucrose synthesis in the day and night converge. Decreased cFBPase expression led to inhibition of sucrose synthesis; accumulation of phosphorylated intermediates; Pi-limitation of photosynthesis; and stimulation of starch synthesis. The starch was degraded to maintain higher levels of sugars and a higher rate of sucrose export during the night. This resembles the response in other species when expression of enzymes in the upper part of the sucrose biosynthesis pathway is reduced. Decreased expression of SPS inhibited sucrose synthesis, but phosphorylated intermediates did not accumulate and carbon partitioning was not redirected towards starch. Sugar levels and sucrose export was decreased during the night as well as during the day. Although ribulose-1,5-bisphosphate regeneration and photosynthesis were inhibited, the PGA/triose-P ratio remained low and the ATP/ADP ratio high, showing that photosynthesis was not limited by the rate at which Pi was recycled during end-product synthesis. Two novel responses counteracted the decrease in SPS expression and explain why phosphorylated intermediates did not accumulate, and why allocation was not altered in the antisense SPS lines. Firstly, a threefold decrease of PPi and a shift of the UDP-glucose/hexose phosphate ratio favoured sucrose synthesis and prevented the accumulation of phosphorylated intermediates. Secondly, there was no increase of AGPase activity relative to cFBPase activity, which would prevent a shift in carbon allocation towards starch synthesis. These responses are presumably triggered when sucrose synthesis is decreased in the night, as well as by day.