Effects of (2 Chloro-Ethyl) Trimethlammonium Chloride on Plant Growth, Leaf Area, and Net Assimilation Rate

Both under glasshouse conditions and in a constant environmentwith light from fluorescent tubes, mustard plants' treatmentwith (2 chloro-ethyl) trimethylammonium chloride (CCC) respondedby an increase in total leaf area. In the glasshouse the increasewas due to the production of more lateral leaves, and in thegrowth chambers to the enlargement of stem leaves, since lateralbranches did not form. Despite the increase in area, the netassimilation rate fell; this may be attributable to an effectof CCC on the photosynthetic mechanism or to an increase inthe mutual shading of leaves, but it is more probable that theinhibition of stem growth also caused by CCC decreases the demandfor photosynthate and leaves photosynthesise less. The leafarea of mustard (a long-day plant) is increased also by short-daytreatment, and here again there is a concomitant shorteningof the stem. Chlorophyll content, both per leaf and per unit area, and totaldry matter per unit area, were increased in tobacco plants grownin culture solutions containing CCC. The treatment affectedthe distribution of nitrogen, increasing the amount per leafand decreasing it per stem.     

Influence of Leaf Starch Concentration on CO(2) Assimilation in Soybean

Net photosynthetic rate, CO₂ compensation concentration, and starch and soluble sugar concentrations were measured in soybean (Glycine max [L.] Merrill) leaves in an attempt to evaluate the effect of carbohydrate concentration on rate of CO₂ assimilation.     

Influence of Flooding on Net CO2Assimilation, Growth and Stem Anatomy of Annona Species

A series of experiments was conducted to assess net CO₂assimilationand growth responses to waterlogging of grafted and seedlingtrees in the genus Annona. Seedlings of A. glabra, A. muricataandA. squamosa L., and scions of ‘Gefner’ atemoya(A. squamosaxA. cherimola Mill.), ‘49-11’ (‘Gefner’atemoyaxA. reticulata L.), ‘4-5’ (‘Priestley’atemoyaxA. reticulata), A. reticulata grafted onto either A.glabra, A. reticulata orA. squamosa rootstocks were floodedfor up to 60 d. Soil anaerobiosis occurred on the third dayof flooding. Seedlings ofA. glabra and A. muricata, and thescions ‘49-11’, ‘Gefner’ atemoya, andA. reticulata grafted onto A. glabra rootstock were consideredflood tolerant based on their ability to survive and grow inflooded conditions. Scions of the normally flood-sensitive A.reticulata, ‘Gefner’ atemoya, and ‘49-11’tolerated root waterlogging when grafted onto the flood-tolerantspecies, A. glabra. In contrast, flooding of A. squamosa seedlingsand rootstocks, and A. reticulata rootstocks greatly reducedgrowth and net CO₂assimilation rates, and resulted in 20–80%tree mortality. Stem anatomical responses to long-term flooding(12 continuous months) were assessed in seedlings of A. glabraand A. muricata, and trees of ‘49-11’ grafted ontoA. glabra. Flooded trees developed hypertrophied stem lenticels,particularly in A. glabra, and enlarged xylem cells resultingin thicker stems with reduced xylem density. Flooding did notincrease air spaces in pre-existing xylem near the pith or inxylem tissue that was formed during flooding. Thus, flood tolerancedid not involve aerenchyma formation in the stem. Copyright1999 Annals of Botany Company Flood tolerance, net CO₂assimilation, photosynthesis, stem anatomy, shoot growth, anaerobiosis, Annonaceae.     

Relationship between Net CO(2) Assimilation and Dry Weight Accumulation in Field-Grown Tobacco

To assess the variability of net photosynthetic CO₂ exchange per unit leaf area and to construct budgets for stands of field-grown tobacco (Nicotiana tabacum, Connecticut Broadleaf), a number of short-time measurements were made on all available leaf positions on two varieties using a hand-held transparent chamber for conducting gas exchange measurements on leaves. Measurements of net CO₂ exchange were carried out on 18 separate days during a 35-day period, beginning 22 days after the seedlings were transplanted to the field. Gas exchange assays on leaves were conducted under ambient conditions of temperature and light intensity at all times of day. Solar radiation was monitored throughout the period, and losses of respiratory CO₂ from stems, roots, and leaves (in the dark) were estimated. A simple model was proposed to relate daily total CO₂ input to irradiance and total leaf area. The total leaf area was assumed to be a function of day number. Dark respiratory losses accounted for 41% to 47% of total CO₂ assimilation. Analysis of variance indicated that the two varieties were not significantly different in whole plant rate of CO₂ fixation per unit of leaf area. CO₂ input was closely associated with leaf area within each variety. Throughout the experiment, the difference between the two varieties in total leaf area per plant was the largest single factor in determining net CO₂ inputs. The cumulative dry weight increase for each variety was similar to the prediction of net dry matter input obtained by gas exchange measurements, thus confirming the close relationship between total plant net CO₂ assimilation and dry weight yield.     

Effects of Ambient and Acute Partial Pressures of Ozone on Leaf Net CO(2) Assimilation of Field-Grown Vitis vinifera L

Mature, field-grown Vitis vinifera L. grapevines grown in open-top chambers were exposed to either charcoal-filtered air or ambient ozone partial pressures throughout the growing season. Individual leaves also were exposed to ozone partial pressures of 0.2, 0.4, or 0.6 micropascals per pascal for 5 hours. No visual ozone damage was found on leaves exposed to any of the treatments. Chronic exposure to ambient O₃ partial pressures reduced net CO₂ assimilation rate (A) between 5 and 13% at various times throughout the season when compared to the filtered treatment. Exposure of leaves to 0.2 micropascals per pascal O₃ for 5 hours had no significant effect on A; however, A was reduced 84% for leaves exposed to 0.6 micropascals per pascal O₃ when compared to the controls after 5 hours. Intercellular CO₂ partial pressure (c_i) was lower for leaves exposed to 0.2 micropascals per pascal O₃ when compared to the controls, while c_i of the leaves treated with 0.6 micropascals per pascal of 0₃ increased during the fumigation. The long-term effects of ambient O₃ and short-term exposure to acute levels of O₃ reduced grape leaf photosynthesis due to a reduction in both stomatal and mesophyll conductances.     

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.     

Acclimation to High CO(2) in Monoecious Cucumbers : I. Vegetative and Reproductive Growth

CO₂ concentrations of 1000 compared to 350 microliters per liter in controlled environment chambers did not increase total fruit weight or number in a monoecious cucumber (Cucumis sativus L. cv Chipper) nor did it increase biomass, leaf area, or relative growth rates beyond the first 16 days after seeding. Average fruit weight was slightly, but not significantly greater in the 1000 microliters per liter CO₂ treatment because fruit numbers were changed more than total weight. Plants grown at 1000 and 350 microliters per liter CO₂ were similar in distribution of dry matter and leaf area between mainstem, axillary, and subaxillary branches. Early flower production was greater in 1000 microliters per liter plants. Subsequent flower numbers were either lower in enriched plants or similar in the two treatments, except for the harvest at fruiting when enriched plants produced many more male flowers than the 350 microliters per liter treatments.     

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.     

Leaf Conductance in Relation to Rate of CO(2) Assimilation: III. Influences of Water Stress and Photoinhibition

Rates of CO₂ assimilation and leaf conductances to CO₂ transfer were measured in plants of Zea mays during a period of 14 days in which the plants were not rewatered, and leaf water potential decreased from −0.5 to −8.0 bar. At any given ambient partial pressure of CO₂, water stress reduced rate of assimilation and leaf conductance similarly, so that intercellular partial pressure of CO₂ remained almost constant. At normal ambient partial pressure of CO₂, the intercellular partial pressure of CO₂ was estimated to be 95 microbars. This is the same as had been estimated in plants of Zea mays grown with various levels of nitrogen supply, phosphate supply and irradiance, and in plants of Zea mays examined at different irradiances.

After leaves of Phaseolus vulgaris L. and Eucalyptus pauciflora Sieb. ex Spreng had been exposed to high irradiance in an atmosphere of CO₂-free N₂ with 10 millibars O₂, rates of assimilation and leaf conductances measured in standard conditions had decreased in similar proportions, so that intercellular partial pressure of CO₂ remained almost unchanged. As the conductance of each epidermis that had not been directly irradiated had declined as much as that in the opposite, irradiated surface it was hypothesized that conductance may have been influenced by photoinhibition within the mesophyll tissue.

     

C(3)-C(4) Intermediate Species in Alternanthera (Amaranthaceae) : Leaf Anatomy, CO(2) Compensation Point, Net CO(2) Exchange and Activities of Photosynthetic Enzymes

Two naturally occurring species of the genus Alternanthera, namely A. ficoides and A. tenella, were identified as C₃-C₄ intermediates based on leaf anatomy, photosynthetic CO₂ compensation point (Γ), O₂ response of г, light intensity response of г, and the activities of key enzymes of photosynthesis. A. ficoides and A. tenella exhibited a less distinct Kranz-like leaf anatomy with substantial accumulation of starch both in mesophyll and bundle sheath cells. Photosynthetic CO₂ compensation points of these two intermediate species at 29°C were much lower than in C₃ plants and ranged from 18 to 22 microliters per liter. Although A. ficoides and A. tenella exhibited similar intermediacy in г, the apparent photorespiratory component of O₂ inhibition in A. ficoides is lower than in A. tenella. The г progressively decreases from 35 microliters per liter at lowest light intensity to 18 microliters per liter at highest light intensity in A. tenella. It was, however, constant in A. ficoides at 20 to 25 microliters per liter between light intensities measured. The rates of net photosynthesis at 21% O₂ and 29°C by A. ficoides and A. tenella were 25 to 28 milligrams CO₂ per square decimeter per hour which are intermediate between values obtained for Tridax procumbens and A. pungens, C₃ and C₄ species, respectively. The activities of key enzymes of C₄ photosynthesis, phosphoenolpyruvate carboxylase, pyruvate Pi dikinase, NAD malic enzyme, NADP malic enzyme and phosphoenolpyruvate carboxykinase in the two intermediates, A. ficoides and A. tenella are very low or insignificant. Results indicated that the relatively low apparent photorespiratory component in these two species is presumably the basis for the C₃-C₄ intermediate photosynthesis.     

Photoreduction of O(2) Primes and Replaces CO(2) Assimilation

A mass spectrometer with a membrane inlet system was used to monitor directly gaseous components in a suspension of algae. Using labeled oxygen, we observed that during the first 20 seconds of illumination after a dark period, when no net O₂ evolution or CO₂ uptake was observed, O₂ evolution was normal but completely compensated by O₂ uptake. Similarly, when CO₂ uptake was totally or partially inhibited, O₂ evolution proceeded at a high (near maximal) rate. Under all conditions, O₂ uptake balanced that fraction of the O₂ evolution which could not be accounted for by CO₂ uptake.     

Leaf Conductance in Relation to Rate of CO(2) Assimilation: I. Influence of Nitrogen Nutrition, Phosphorus Nutrition, Photon Flux Density, and Ambient Partial Pressure of CO(2) during Ontogeny

Plants of Zea mays were grown with different concentrations of nitrate (0.6, 4, 12, and 24 millimolar) and phosphate (0.04, 0.13, 0.53, and 1.33 millimolar) supplied to the roots, photon flux densities (0.12, 0.5, and 2 millimoles per square meter per second), and ambient partial pressures of CO₂ (305 and 610 microbars). Differences in mineral nutrition and irradiance led to a large variation in rate of CO₂ assimilation per unit leaf area (A, 11 to 58 micromoles per square meter per second) when measured under standard conditions. The variation was shown, with the plants that had received different amounts of nitrate, to be related to variations in the nitrogen and chlorophyll contents, and phosphoenolpyruvate and ribulose-1,5-bisphosphate carboxylase activities per unit leaf area. Irrespective of growth treatment, A and leaf conductance to CO₂ transfer (g), measured under standard conditions were in almost constant proportion, implying that intercellular partial pressure of CO₂ (p_i), was almost constant at 95 microbars. The same proportionality was maintained as A and g increased in an initially nitrogen-deficient plant that had been supplied with abundant nitrate. It was shown that p_i measured at a given ambient partial pressure was not affected by the ambient partial pressure at which the plants had been grown, although it was different when measured at different ambient partial pressures. This suggests that the close coupling between A and g in these experiments is not associated with sensitivity of stomata to change in p_i.

Similar, though less comprehensive, experiments were done with Gossypium hirsutum, and yielded similar conclusions, except that the proportionality between A and g at normal ambient partial pressure of CO₂ implied P_i ≈ 200 microbars.

     

Interrelations of Flowering and Vegetative Growth in Callistephus chinensis (var. Queen of the Market)

In a study of the effect of photoperiod upon the growth andflowering of Calistephus chinensis (var. Queen of the Market)it has been shown that a one-hour night break of low intensityred light given to plants growing in eight hours daylight canhave a number of morphogenetic effects. In the young plant, such a treatment increased the total areaof the leaf surface and the area per unit weight of leaf material,i.e. the specific leaf area. This effect enabled the treatedplants to make greater use of the incident light, for afternine weeks they were at least 25 per cent heavier than comparableplants which had not had the benefit of the night-break treatment.This treatment also caused flower induction and concomitantstem extension, but transfer back into eight-hour days afteran inductive period accelerated further flower development andrestricted stem extension of both the main axis and the laterals.If flower development was delayed by continuing night breaksor by delaying the onset of induction then more flowers wereeventually formed, but in the very prolific treatments flowersize was reduced. The experiment also indicated that the partition of availabledry weight between leaves, stems, and roots followed a definitepattern dependent only upon total plant dry weight. The diversionof dry weight into flowers was strongly accelerated by transferinto eight-hour days after induction but the remaining dry matterstill appeared to be distributed between the vegetative partsalong the general pattern determined by total vegetative weight. Finally the experiment showed that a wide range of plant formsbearing varying amounts of flowers could be obtained by appropriatetransfers between treatments.     

Net CO2 Exchange Rate and Shoot Growth of Young Poplar (Populus) Clones

Eighteen poplar (Populus) clones, varying in growth capacitywere grown in plastic containers under outdoor conditions. Duringthe first year of growth their net CO₂ exchange rate (NCER)was studied by infra-red gas analysis as a function of photosyntheticphoton flux density (PPFD) under controlled environment conditions.Maximal NCER (under saturating PPFD) and the PPFD compensationpoint were significantly correlated with the first year's shootlength and hence with the above ground biomass production ofthe various clones. Key words: Poplar, Shoot growth, CO₂ exchange rate     

Acclimation of CO(2) Assimilation in Cotton Leaves to Water Stress and Salinity

Cotton (Gossypium hirsutum L. cv Acala SJ2) plants were exposed to three levels of osmotic or matric potentials. The first was obtained by salt and the latter by withholding irrigation water. Plants were acclimated to the two stress types by reducing the rate of stress development by a factor of 4 to 7. CO₂ assimilation was then determined on acclimated and nonacclimated plants. The decrease of CO₂ assimilation in salinity-exposed plants was significantly less in acclimated as compared with nonacclimated plants. Such a difference was not found under water stress at ambient CO₂ partial pressure. The slopes of net CO₂ assimilation versus intercellular CO₂ partial pressure, for the initial linear portion of this relationship, were increased in plants acclimated to salinity of −0.3 and −0.6 megapascal but not in nonacclimated plants. In plants acclimated to water stress, this change in slopes was not significant. Leaf osmotic potential was reduced much more in acclimated than in nonacclimated plants, resulting in turgor maintenance even at −0.9 megapascal. In nonacclimated plants, turgor pressure reached zero at approximately −0.5 megapascal. The accumulation of Cl⁻ and Na⁺ in the salinity-acclimated plants fully accounted for the decrease in leaf osmotic potential. The rise in concentration of organic solutes comprised only 5% of the total increase in solutes in salinity-acclimated and 10 to 20% in water-stress-acclimated plants. This acclimation was interpreted in light of the higher protein content per unit leaf area and the enhanced ribulose bisphosphate carboxylase activity. At saturating CO₂ partial pressure, the declined inhibition in CO₂ assimilation of stress-acclimated plants was found for both salinity and water stress.     

Effect of Growth Regulators on CO(2) Assimilation in Leaves, and its Correlation with the Bud Break Response in Photosynthesis

Experiments have been done to confirm the previously reported effect of indoleacetic acid (IAA) on the rate of CO₂ assimilation in bean leaves. It was shown that spraying the leaves of a variety of plants caused an increase in the rate of CO₂ assimilation from 30% to 100% during the half-hour to 1 hour period following spraying. The only plant tested which did not show such an effect was corn.

The breaking of dormancy of axial buds in the bean plant was correlated with an increase in the rate of CO₂ assimilation in adjacent leaves for a brief period of time. It has been shown that IAA solution sprayed on 1 leaflet of a leaf can cause an increase in the rate of CO₂ assimilation in the other leaflets, and that IAA applied to the cut stem of a leaflet or a developing bud can be transported to adjacent leaves and cause an increase in the CO₂ assimilation rate. The reaction caused by IAA is very similar to that caused by the breaking of dormancy of a bud. This indicates that the bud break response in CO₂ assimilation in leaves is caused by auxin synthesized in a bud as it begins to grow, and exported into adjacent leaves.

     

Leaf Conductance in Relation to Rate of CO(2) Assimilation: II. Effects of Short-Term Exposures to Different Photon Flux Densities

When photon flux density incident on attached leaves of Zea mays L. was varied from the equivalent of 0.12 of full sunlight to full sunlight, leaf conductance to CO₂ transfer, g, changed in proportion to the change in rate of CO₂, assimilation, A, with the result that intercellular partial pressure of CO₂ remained almost constant. The proportionality was the same as that previously found in g and A measured at one photon flux density in plants of Zea mays L. grown at different levels of mineral nutrition, light intensities, and ambient partial pressures of CO₂. In shade-grown Phaseolus vulgaris L. plants, A as photon flux density was increased from about 0.12 up to about 0.5 full sunlight, the proportionality being almost the same in plants grown at low and at high light intensity.

When photon flux density incident on the adaxial and abaxial surfaces of the isolateral leaves of Eucalyptus pauciflora Sieb. ex Spreng was varied, g and A also varied proportionally. The leaf conductance in a particular surface was affected by the photon flux density at the opposite surface to a greater extent than was expected on the basis of transmittance. The results indicated that stomata may, in some way, be sensitive to the photon flux absorbed within the leaf as a whole.

     

Contribution of Metabolites of Photosynthesis to Postillumination CO(2) Assimilation in Response to Lightflects

In the shade plant Alocasia macrorrhiza grown in low light, photosynthetic CO₂ assimilation during a 5 second lightfleck plus postillumination CO₂ assimilation can allow up to 60% more photosynthesis than that which occurs during 5 seconds of steady state light of the same intensity (RL Chazdon, RW Pearcy 1986 Oecologia. 69: 524-531). Metabolites of photosynthesis were measured to determine if the pool of ribulose 1,5-bisphosphate (RuBP) could account for all of the postillumination CO₂ assimilation following a lightfleck in Alocasia. It was found that the pool of triose-P was much larger than that of RuBP and could account for five times more postillumination CO₂ assimilation than could RuBP. The same trend was seen in the sun plant Phaseolus vulgaris when it was grown in the shade. In contrast, sun-grown Alocasia and Phasiolus did not have a large pool of triose-P relative to RuBP following a lightfleck. In sun plants, carbon may rapidly be converted to RuBP in the light whereas in shade plants there may be a restriction in the path between the triose-P and RuBP pools. It is hypothesized that in shade plants the buildup of triose-P rather than RuBP during the lightfleck prevents inhibition of electron transport which may otherwise occur because of competition for ATP between the two kinases of the photosynthetic carbon reduction cycle. Utilization of the triose-P for postillumination CO₂ fixation would require the capacity for significant postillumination ATP synthesis. The extensive grana stacking and large intrathylakoid space which accompanies the high level of chlorophyll in low-light-grown Alocasia could be an important contributing factor to postillumination ATP formation.     

Effect of Triacontanol on Chlamydomonas: I. Stimulation of Growth and Photosynthetic CO(2) Assimilation

Treatment of Chlamydomonas reinhardtii cells, cultured at 5% CO₂, with 1 to 1000 micrograms triacontanol (TRIA) per liter resulted in 21 to 35% increases in cell density, 7 to 31% increases in total chlorophyll, and 20 to 100% increases in photosynthetic CO₂ assimilation. The increase in CO₂ fixation with TRIA treatment occurred before, and was independent of, increases in total chlorophyll or cell number. Chlamydomonas cells responded to a broad range of TRIA concentrations that were at least one order of magnitude above the optimum concentration established for higher plants. The necessity for larger concentrations of TRIA may be due to destabilizing effects of Ca²⁺ and K⁺ present in the Chlamydomonas growth medium. These ions caused flocculation of the colloidally dispersed TRIA in apparent competition with binding of [¹⁴C]TRIA to Chlamydomonas cells. Octacosanol inhibited the effect of TRIA on photosynthetic CO₂ assimilation. TRIA treatment did not alter the distribution of ¹⁴C-label among photosynthetic products. The effect of TRIA on photosynthetic CO₂ assimilation increased with time after treatment up to 3 days. Chlamydomonas cells that had been grown at low-CO₂ (air) did not respond to TRIA, and transfer of high-CO₂ (5%) grown cells that had responded to TRIA to a low-CO₂ atmosphere resulted in a loss of the effect of TRIA. The effect of pH on photosynthetic CO₂ assimilation indicated that CO₂ is probably the species of inorganic carbon utilized by control and TRIA-treated Chlamydomonas cells.