I. S. Drozdova

LIGHT RESPONSE CURVES OF PHOTOSYNTHESIS IN LEAVES OF SUN-TYPE AND SHADE-TYPE PLANTS GROWN IN BLUE OR RED LIGHT

Abstract Light saturation curves after growth in blue light (BL) or red light (RL) were studied in leaves of the shade-type C 3 -plant, Convallaria majalis , and two sun-type plant species, the C 3 -plant, Hordeum vulgare , and the C 4 -plant, Zea mays . The photosynthetic activity of BL leaves of C. majalis was found to be 1.5 times higher than that of RL leaves at all the irradiances examined. Unlike the shade-type plant, BL promoted photosynthesis in the leaves of the two sun-type plants more strongly for saturating irradiance levels (activities 1.4–1.5 times higher than in RL) than for the linear part of the light saturation curve (1.1–1.2 times higher than in RL). The results indicate that the enhancing action of BL on the different parts of the photosynthetic apparatus depends on the ecological type of plant: in contrast with shade-type plants, BL promotes mainly the activity of dark photosynthetic reactions in leaves of sun-type plants.

Endogenous Control of Photosynthetic Activity during Progressive Drought: Influence of Final Products of Photosynthesis

Photosynthetic activities and the redox states of photosystem I (PSI) and photosystem II (PSII) in intact leaves of cucumber plants (Cucumis sativus L.), as well as the sucrose and starch contents were examined under conditions of ongoing soil water deficit imposed by the cessation of watering. As the soil drought progressed, the maximum rate of photosynthetic CO2 fixation was shown to decrease. These changes in the maximum photosynthetic rate occurred synchronously with changes in the maximum quantum yield of photosynthesis. Under soil water deficit, the reduced form of PSII primary acceptor QA was accumulated only at photon flux densities of about 100 μmol/(m2 s). At such photon flux densities, the changes in nonphotochemical quenching (qN) induced by soil water deficit were opposite to changes in photochemical quenching parameter (1 – qP). Irrespective of the duration of soil drought, the relationship between steady-state concentrations of photochemically inactive reaction centers of PSI and PSII (the fractions of P700 and QA in the oxidized and reduced state, respectively) was almost linear, which provides evidence for the concerted operation of both photosystems. The conditions of soil water deficit promoted sucrose accumulation in the source leaf, which was paralleled by a substantial decrease in the amount of starch in the same leaf. The highest content of sucrose in the leaf after a 7-day drought was correlated with the largest decrease in photosynthetic activity. It is concluded that the progressive drought triggers an endogenous mechanism that regulates photosynthesis through feedback relations, namely, the inhibition of photosynthesis by its end products.

Effects of Light Spectral Quality on Morphogenesis and Source–Sink Relations in Radish Plants

The accumulation of dry matter and the content of major phytohormones in the aboveground and underground plant parts, as well as light curves and the diurnal course of photosynthesis in the leaves were studied in radish (Raphanus sativusL.) plants of different ages that were grown under red (RL) or blue (BL) light. As seen from the rapid increase in plant biomass, the development of storage organs (hypocotyl or tap root) started on the 14th day after the emergence of seedling of the BL plants and on the 21st day for the RL plants. Conversely, RL stimulated biomass accumulation in the aboveground parts (petioles and stems) already in the early stages of plant development. Light spectral quality only slightly affected the activity and the diurnal course of photosynthesis. The GA content was ten times higher in the aboveground parts of the RL plants than those of the BL plants. The hypocotyl of the BL plants contained much higher amounts of cytokinins and IAA than that of the RL plants. The specific responses of the source–sink relations to the light quality were related to the distribution of various phytohormones between the aboveground and underground parts of the plants: RL increased the content of gibberellins (GA) in the aboveground parts of plants, thus increasing their sink activity, whereas BL stimulated the synthesis of cytokinins and IAA in the hypocotyl and enhanced its development. Light quality-specific morphogenetic responses were reversed when plants were treated with exogenous GA or paclobutrazol, an inhibitor of GA synthesis. The treatment of the BL plants with exogenous GA stimulated petiole and hypocotyl elongation and induced stem formation. The treatment of the BL plants with paclobutrazol led to shortened petioles, the flattening of the storage organ, and the disappearance of the stem.

Leaf Growth, Photosynthetic Rate, and Phytohormone Contents in Cucumis sativus Plants under Progressive Soil Drought

Effects of progressive soil drought on leaf growth, the rate of photosynthesis, and phytohormone contents were followed in the experiments with cucumber (Cucumis sativus L.) plants. Suppression of photosynthesis by drought did not immediately cause growth retardation, because the latter was observed one day earlier than the inhibition of photosynthesis. In the meantime, growth retardation could be caused by a decline in IAA and cytokinin contents, rather than ABA accumulation, because ABA accumulated when the growth has been already suppressed.

Photosynthesis and Dark Respiration in Leaves of Different Ages of Partly Flooded Maize Seedlings

Maize seedlings were flooded for periods from 1 to 15 days, and the leaves of different ages were then taken to examine photosynthesis, dark respiration, transpiration, chlorophyll content, and some morphometric parameters. The responses of leaves to root submergence essentially depended on the leaf layer and the treatment duration. A short-term flooding (1–24 h) induced primary stress responses in the first leaf. Photosynthesis and respiration in this leaf oscillated around the control levels with amplitudes of ±15–25% and ±40–60%, respectively. After a longer flooding, the CO2 exchange in the second leaf was suppressed, while oxygen uptake was stimulated. In the third leaf, which was formed during submergence, the photosynthetic rate increased and the respiratory activity decreased. The transpiration rate did not change in these leaves for 15 days of flooding. The hypoxic treatment, at its early stages, retarded growth and disturbed the source–sink relations. At later stages the plants adapted to hypoxic environment: the seedling growth was restored, which elevated the demand for assimilates and stimulated photosynthesis. It is concluded that plants overcome negative impact of the root hypoxia at the systemic level.

Long-term effects of blue or red light on ATP and ADP contents in primary barley leaves

Levels of ATP and ADP were studied in primary leaves of barley (Hordeum vulgare L. cv. Viner) seedlings grown under blue (BL) or red light (RL) of various irradiances. In mature leaf segments, BL stimulated a greater accumulation of adenylates than RL. Transfer of barley seedlings from RL to BL for 48 h caused about a twofold increase in the content of adenylates, probably due to de-novo synthesis of adenine nucleotides. Weak BL was found to stimulate an increase in the adenylate content and a higher irradiance enhanced the stimulatory effect. The adenylate content increased markedly from the base towards the tip of barley leaves grown under BL but not in those grown under RL. However, the adenylate content was higher in the basalmost segment of barley leaves grown under RL, indicating that the action of RL on adenylate content proceeded more rapidly than that of BL. The same conclusion could be drawn from the results of experiments with de-etiolated leaves. A linear relationship was established between the maximum rate of CO2 fixation and the ATP or ADP content in mature segments of primary barley leaves. The possible involvement of two photoreceptors, phytochrome and cryptochrome, in the long-term light regulation of the total content of adenylates in primary barley leaves is discussed.

Modulating Effect of Far-Red Light on Activities of Alternative Electron Transport Pathways Related to Photosystem I

Barley (Hordeum vulgare L.) leaves were irradiated with far-red (FR) light of various intensities after different periods of dark adaptation in order to investigate activities of alternative electron transport pathways related to photosystem I (PSI). Photooxidation of P700, the primary electron donor of PSI, was saturated at FR light intensity of 0.15 µmol quanta/(m2 s). As the photon flux density was raised in this range, the slow and middle components in the kinetics of P700+ dark reduction increased, whereas the fast component remained indiscernible. The amplitudes of the slow and middle components diminished upon further increase of FR photon flux density in the range 0.15–0.35 µmol quanta/(m2 s) and remained constant at higher intensities. The fast component of P700+ reduction was only detected after FR irradiation with intensities above 0.15 µmol quanta/(m2 s); the light-response curve for this component was clearly sigmoid. In dark-adapted barley leaves, three stages were distinguished in the kinetics of P700 photooxidation, with the steady state for P700+ achieved within about 3 min. In leaves predarkened for a short time, the onset of FR irradiation produced a very rapid photooxidation of P700. As the duration of dark exposure was prolonged, the amplitude of the first peak in the kinetic curve of photoinduced P700 photooxidation was diminished and the time for attaining the steady-state oxidation level was shortened. After a brief dark adaptation of leaves, ferredoxin-dependent electron flow did not appreciably contributed to the kinetics of P700+ dark reduction, whereas the components related to electron donation from stromal reductants were strongly retarded. It is concluded that FR light irradiation, selectively exciting PSI, suffices to modulate activities of alternative electron transport routes; this modulation reflects the depletion of stromal reductants due to continuous efflux of electrons from PSI to oxygen under the action of FR light.

Temporal Pattern of Photosynthesis under Continuous Illumination of Radish Plants

Changes in photosynthetic activity, redox state of photosystem I (PSI) and photosystem II (PSII), as well as starch and sucrose content were studied on the source leaves of 18- to 20-day-old radish (Raphanus sativus L.) plants that were dark-adapted for 12 h and then exposed to continuous white light (170 μmol quanta/(m2 s)). The kinetic pattern of photosynthetic activity comprised three phases. Within the first 6 h of light adaptation (first phase), the maximum photosynthetic rate and the quantum yield of photosynthesis increased 1.6 times in the illuminated leaves compared to the leaves of plants placed in darkness. Further illumination led to the decrease of both photosynthetic indices by about 20% (12 h after the onset of light exposure, second phase) and finally increased them to the level observed after 6-h light exposure (72 h, third phase). The stationary photooxidation level of PSI primary donor was relatively low within the first 6 h of light adaptation, and then it steeply increased. The linear relationship between the amounts of photoreduced PSII primary acceptor and photooxidized PSI primary donor did not change during prolonged light adaptation, showing a highly coordinated functioning of both photosystems. The amount of sucrose in leaves attained its peak after 12 h of light adaptation and did not change further on. The starch content increased to its peak within 24 h of illumination and decreased gradually upon longer exposures. It is concluded that, despite active export of assimilates to the developing storage organ, the source leaves exhibit a nonmonotonic temporal course of endogenously regulated photosynthetic activity, which was related to changes in the effectiveness and, possibly, the number of the components of photosynthetic apparatus.

Distribution of Phytohormones among Different Plant Organs Determines the Development of Competitive Sinks in Radish Plants

Phytohormones are known to mediate many responses of plants to changes in environmental cues including light climate. Light quality governs various stages of plant growth and development (Short and Briggs, 1994; Smith, 1995). One of the most important features of plant photoregulation is involvement of light quality in the determination of sink strength (Aksenova et al., 1994). Several sinks, such as newly developing leaves, petioles and stem, and storage organ(s) compete for photosynthetic metabolites in a plant. Some plant organs dominate others in their ability to attract assimilates from donor leaves, depending on their sink strength (Drozdova et al., 1987). Correspondingly, various hormones have been proposed to stimulate metabolite flow to above- and/or underground plant organs (Palmer and Smith, 1969; Metzger, 1988). It is possible therefore, that light quality changes the direction of assimilate flow in a plant through alterations in hormonal status of plant organs. We show here that hormones belonging to different classes (gibberellins, cytokinins, and auxin) act in concert to determine preferential development of either above-ground sink (petioles and stem) under red light (RL) or underground sink (storage organ) under blue light (BL).