Carbon gain and photosynthetic response of chrysanthemum to photosynthetic photon flux density cycles

Most models of carbon gain as a function of photosynthetic irradiance assume an instantaneous response to increases and decreases in irradiance. High- and low-light-grown plants differ, however, in the time required to adjust to increases and decreases in irradiance. In this study the response to a series of increases and decreases in irradiance was observed in Chrysanthemum × morifolium Ramat. “Fiesta” and compared with calculated values assuming an instantaneous response. There were significant differences between high- and low-light-grown plants in their photosynthetic response to four sequential photosynthetic photon flux density (PPFD) cycles consisting of 5-minute exposures to 200 and 400 micromoles per square meter per second (μmol m⁻²s⁻¹). The CO₂ assimilation rate of high-light-grown plants at the cycle peak increased throughout the PPFD sequence, but the rate of increase was similar to the increase in CO₂ assimilation rate observed under continuous high-light conditions. Low-light leaves showed more variability in their response to light cycles with no significant increase in CO₂ assimilation rate at the cycle peak during sequential cycles. Carbon gain and deviations from actual values (percentage carbon gain over- or underestimation) based on assumptions of instantaneous response were compared under continuous and cyclic light conditions. The percentage carbon gain overestimation depended on the PPFD step size and growth light level of the leaf. When leaves were exposed to a large PPFD increase, the carbon gain was overestimated by 16 to 26%. The photosynthetic response to 100 μmol m⁻² s⁻¹ PPFD increases and decreases was rapid, and the small overestimation of the predicted carbon gain, observed during photosynthetic induction, was almost entirely negated by the carbon gain underestimation observed after a decrease. If the PPFD cycle was 200 or 400 μmol m⁻² s⁻¹, high- and low-light leaves showed a carbon gain overestimation of 25% that was not negated by the underestimation observed after a light decrease. When leaves were exposed to sequential PPFD cycles (200-400 μmol m⁻² s⁻¹), carbon gain did not differ from leaves exposed to a single PPFD cycle of identical irradiance integral that had the same step size (200-400-200 μmol m⁻² s⁻¹) or mean irradiance (200-300-200 μmol m⁻² s⁻¹).     

Sequential response of whole plant water relations to prolonged soil drying and the involvement of xylem sap ABA in the regulation of stomatal behaviour of sunflower plants

Sunflower plants ( Helianihus animus cv. Tall Single Yellow} were grown in the greenhouse in drain pipes (100 mm inside diameter and 1 m long) rilled with John Innes No. 2 compost. When the fifth leaf had emerged, half of the plants were left unwatered for 6 days, rewatered for 2 days and then not watered for another 12 days. Measurements of water relations and abaxial stomatal conductance were made at each leaf position at regular intervals during the experimental period. Estimates were also made of soil water potentials along the soil profile and of ABA concentrations in xylem sap and leaves.
Soil drying led to some reduction in stomatal conductance alter only 3 days but leaf turgors were not reduced until day 13 (6 days after rewatering). When the water relations of leaves did change, older leases became substantially dehydrated while high turgors were recorded in younger leaves. Leaf ABA content measured on the third youngest leaf hardly changed over the first 13 days of the experiment, despite substantial soil drying, while xylem ABA concentrations changed very significantly and dynamically as soil water status varied, even when there was no effect of soil drying on leaf water relations. We argue that the highest ABA concentrations in the xylem, found as a result of substantial soil drying, arise from synthesis in both the roots and the older leaves, and act to delay the development of water deficit in younger leases.
In other experiments ABA solutions were watered on to the root systems of sunflower plants to increase ABA concentrations in xylem sap. The stomatal response to applied ABA was quantitatively very similar to that to ABA generated as a result of soil drying. There was a log-linear relationship between the reduction of leaf conductance and the increase of ABA concentration m xylem sap.     

A feedback control system for the precise and continuous regulation of photosynthetic photon flux density

A simple and inexpensive feedback control system that provides continuous and precise control of photosynthetic photon flux density (PPFD) in a whole plant cuvette is described. A ‘Plexiglass’ tank is interposed between a light source and cuvette and PPFD changed by varying the level of dyed liquid in the tank. The amount of liquid pumped into or drained from the tank is a function of the difference (error) between a defined set point value of PPFD and that measured in the cuvette. The set point can be varied as a function of time, can follow the output of a quantum sensor measuring ambient PPFD or can be driven by values of PPFD read from a data file. Within the 0.4 to 0.64 μm waveband, the dye acts as a neutral density filter so that there is no change in spectral distribution with PPFD. Photosynthetic photon flux density in the cuvette was controlled to better than 20 μmol m⁻²s⁻¹ when the set point was varied from 200 to 1100 μmol m⁻²s⁻¹ over 3 min. When the set point was held constant or changed less rapidly, errors did not exceed 5 μmol m⁻²s⁻¹. Net photosynthesis of Western redcedar (Thuja plicata Donn.) seedlings held at 18 °C closely followed rapid changes in PPFD.     

Responses of photosynthesis and carbohydrate-partitioning to limitations in nitrogen and water availability in field-grown sunflower*

Abstract. Sunflower plants (Helianthus annuus L., cv. CGL 208) were field-grown in adjacent plots of varying resource availability. Control plants received irrigation (on a 4–5 d interval) and high levels of fertilizer nitrogen. Nutrient-stress (N-stress) plants received control levels of irrigation but no nutrient amendments and were determined to be nitrogen-limited. Water-stress (H₂O-stress) plants received control levels of fertilizer nitrogen, but no irrigation after approximately 6 weeks of plant growth. Both stress treatments reduced maximum and diurnal net photosynthesis (A) but resulted in different physiological or biochemical adjustments that tended to maintain or increase A per unit of resource (nitrogen or water) in shortest supply while decreasing the ratio of A per unit of abundant resource. Nutrient-stress reduced total foliar nitrogen, foliar chlorophyll, and initial and total RuBPCase activities, thereby enhancing or preserving photosynthetic nitrogen-use efficiency (NUE), defined as the maximum A observed per unit of leaf nitrogen, relative to the control and H₂O-stress treatments. In addition, N-stress reduced photosynthetic water-use efficiency (WUE), defined as the ratio of A to stomatal conductance to water vapour (g). The slope of A versus g increased with H₂O-stress. In addition, sunflower plants responded to H₂O-stress by accumulating foliar glucose and sucrose and by exhibiting diurnal leaf wilting, which presumably provided additional improvements in photosynthetic WUE through osmoregulation and reduction of midday radiation interception respectively. Photosynthetic NUE was decreased by H₂O-stress in that control levels of total nitrogen, foliar chlorophyll, and RuBPCase activities were maintained even after mean diurnal levels of A had fallen to less than 50% of the control level. We conclude that field-grown sunflower manages a trade-off between photosynthetic WUE and NUE, increasing use efficiency of the scarce resource while decreasing use efficiency of the abundant resource.     

Regulation of the photosynthetic capacity of primary bean leaves by the red:far-red ratio and photosynthetic photon flux density of incident light

The main objective of the present work was to examine the effects of the red:far-red ratio (R:FR) prevailing during leaf development on the photosynthetic capacity of mature leaves. Plants of Phaseolus vulgaris L. cv. Balin de Albenga were grown from time of emergence in a controlled environment room, 25 ± 3°C, 12-h photoperiod, with different light treatments:a) high photosynthetic photon flux density (PPFD) = 800 μmol m⁻¹ s⁻¹+ high R:FR= 1.3;b) low PPFD= 300 μmol m⁻² s⁻¹+ high R:FR= 1.3; c) high PPFD=800 μmol m⁻² s⁻¹+ low R:FR= 0.7; d) low PPFD= 300 μmol m⁻²s⁻¹+ low R:FR=0.7. With an R:FR ratio of 1.3, a decrease in irradiance during leaf growth reduced photosynthesis when measured at moderate to high PPFD; but when measured at low PPFD, leaves expanded under low irradiance actually had photosynthesis rates higher than those of leaves grown in high irradiance. A low R:FR ratio during development reduced the photosynthetic capacity of the leaves. In leaves expanded under R:FR = 0.7 and high irradiance photosynthesis was reduced by 42 to 89%, depending on the PPFD at which measurements were made, whereas for leaves developed at R:FR = 0.7 and low irradiance photosynthesis decreased by 21 to 24%, compared to leaves under R:FR = 1.3 and similar irradiance. The reduced photosynthetic capacity under R:FR = 0.7 and high irradiance. In natural environments, leaves may experience low R:FR conditions temporarily during their development, and this may affect their future photosynthetic capacity in full sunlight.     

红松阔叶混交林林隙光量子通量密度的时空分布格局

以小兴安岭原始红松阔叶混交林林隙为对象,采用网格法布点,对生长季林隙内各样点光量子通量密度(photosynthetic photon flux density,PPFD)进行连续观测,利用基本统计学和地统计学方法分析其时空分布格局.结果表明:红松阔叶混交林林隙的PPFD高值区日变化明显,最大值出现在12:00,位于林隙北侧.林隙的PPFD 6月最高,7、8、9月依次递减,其中7月PPFD不同位置间的变异系数最大;各月均为中等变异.不同月份林隙PPFD空间异质性的强度和尺度不同,6月变程最大,7月基台值和结构比最大;各月林隙PPFD斑块复杂程度不同,最大值均位于林隙东北侧.郁闭林分和空旷地的月平均PPFD变化次序与林隙一致.各样点月平均PPFD为空旷地最高,林隙次之,郁闭林分最低.     

Spatial heterogeneity of photosynthetic photon flux density in the canopy ofMiscanthus sinensis

Spatial variation in photosynthetic photon flux density (PPFD) was investigated in detail at different heights within the canopy of aMiscanthus sinensis grassland to evaluate the light environment of microsites for establishment of heliophilic tree seedlings. Highly heterogeneous patterns of light distribution were revealed within the apparently uniform grass canopies, especially under direct light. The frequency distribution patterns of relative PPFD (RPFD) were compared among different solar and sky conditions. With increasing height in the canopy, the mean RPFD value and standard deviation (SD) increased, while the skewness and kurtosis of the distribution decreased. The mean RPFD and SD were higher, especially at higher solar elevation angles, under direct light than those under diffuse light conditions. The frequency distribution of RPFD was more platykurtic under direct light and at higher solar elevation angles.     

Early degradation of photosynthetic membranes in carob and sunflower cotyledons

The assimilatory activity of cotyledons can play an essential role in the survival of seedlings with a slow and delayed development of primary leaves. Changes in the photosynthetic activity of the cotyledon, from the onset of greening through senescence, were studied in two such plants, carob and sunflower, in order to determine its efficiency and duration, also in connection with the achievement of assimilatory autonomy by the plantlet. Chlorophyll analyses showed that the cotyledon's chloroplasts reached maximal greening in plantlets with a pair of expanded leaves. In contrast, the cotyledon's photosynthetic activity, measured as the rate of oxygen release, started to decrease early, before expansion of primary leaves. The decrease was due to the inactivation of a number of photosystem II (PSII) units, as revealed by immunodetection of breackdown products of the reaction centre's D1 and D2 thylakoid proteins. No signals of PSII alteration were noticed in the primary leaf chloroplasts that differentiated under the same environmental conditions. The damage to the cotyledon PSII, occurring in a non-photoinhibitory situation, might be due to a slower rate of turnover of D1 polypeptide than in the leaf thylakoids. The differential turnover of this protein in cotyledons and in leaves might represent an organ-specific regulation of the photosynthetic activity. The peculiarity of the cotyledon thylakoids make these organs useful objects for studying the metabolic cycle of both D1 and D2 proteins in vivo, under non-photoinhibiting conditions.     

Etiolation symptoms in sunflower (Helianthus annuus) cotyledons partially covered by the pericarp of the achene

Background and Aims

Etiolation symptoms and the greening process are usually studied on dark-germinated seedlings and this raises the question – can these results be generalized for plants growing under field conditions? This work examines various aspects of the plastid differentiation under the covering of the achene wall, which often remains attached to the cotyledons of sunflower (Helianthus annuus) seedlings grown under light.

Methods

Cotyledons of 7- to 10-d-old sunflower seedlings grown in the dark and on light were examined. The partially covered cotyledons were sectioned into light-exposed, covered and transition zones. Pigment contents, 77 K fluorescence spectroscopy, electron microscopy and fluorescence imaging, along with fluorescence kinetic methods, were used.

Key Results

The light-exposed zone of the partially covered cotyledons was similar to cotyledons developed without achene covering. However, some of the plastids had prolamellar bodies among the granal thylakoid membranes; despite this no protochlorophyllide was detected. The fully covered, yellowish sections contained protochlorophyllide forms emitting at 633 and 655 nm and well-developed prolamellar bodies, similar to those of etiolated cotyledons. In addition, reduced amounts of chlorophyll a, chlorophyll b and stacked thylakoid membrane pairs were found in this region. The transitional sections showed a mixture of the characteristics of the covered and exposed sections. Various, but significantly different values of the photosynthetic activity parameters were found in each sector of the partially covered cotyledons.

Conclusions

The partial covering of the achene wall shades the cotyledon tissues effectively, enough to provoke the appearance of etiolation phenomena, i.e. the permanent presence of flash-photoactive protochlorophyllide complexes and prolamellar bodies (with or without protochlorophyllide), which proves that these phenomena may appear under natural illumination conditions.Key words: Cotyledon, etio-chloroplast, etioplast, etiolation, Helianthus annuus, photosynthetic activity, protochlorophyllide, prolamellar body, sunflower     

Leaf photosynthetic and solar-tracking responses of mallow, Malva parviflora, to photon flux density

Malva parviflora L. (mallow) is a species that occupies high-light habitats as a weedy invader in orchards and vineyards. Species of the Malvaceae are known to solar track and anecdotal evidence suggests this species may also. How M. parviflora responds physiologically to light in comparison with other species within the Malvaceae remains unknown. Tracking and photosynthetic responses to photon flux density (PFD) were evaluated on plants grown in greenhouse conditions. Tracking ability was assessed in the growth conditions and by exposing leaves to specific light intensities and measuring changes in the angle of the leaf plane. Light responses were also determined by photosynthesis and chlorophyll fluorescence. Leaves followed a heliotropic response which was highly PFD-dependent, with tracking rates increasing in a curvilinear pattern. Maximum tracking rates were up to 20°h⁻¹ and saturated for light above 1300 μmol (photons) m⁻² s⁻¹. This high-light saturation, both for tracking (much higher than the other species), and for photosynthesis, confirmed mallow as a high-light demanding species. Further, because there was no photoinhibition, the leaves could capture the potential of an increased carbon gain in higher irradiance by resorting to solar tracking. Modelling suggested the tracking response could increase the annual carbon gain by as much as 25% compared with leaves that do not track the sun. The various leaf attributes associated with solar tracking, therefore, help to account for the success of this species as a weed in many locations worldwide.     

The effects of illumination sequence, CO2 concentration, temperature and acclimation on the convexity of the photosynthetic light response curve

It was shown previously that the convexity (curvature or rate of bending) of the photosynthetic light response curve was strongly correlated with chlorophyll content in shade acclimated conifer needles (Leverenz 1987, Physiol. Plant. 71: 20–29), in agreement with an hypothesis that gradients of light within leaves affect the convexity. In the present study it is shown that the convexity at any given chlorophyll content can be altered when leaves of Pinus sylvestris L. Picea glauca (Muench), Picea mariana (M.II.) BS.P. and Picea abies (L.) Karst pre-treated with less shade. This probably induced a differential acclimation of cells on the top and bottom sides of the leaves to their local light environment. Leaves were illuminated on i) their top surface, ii) their bottom surface, or iii) uniformly in a light integrating sphere during measurements of photosynthesis. After shoots had been transferred from their growth environment to a new measuring environment, the convexity increased from the first to the second day towards a maximum of 0.97. The rate of increase towards this maximum was 55 to 62% per day and probably is the result of re-acclimation of cells within the leaves. The data shown that the act of measuring photosynthesis induces a significant alteration in the experimental material when measurements are made for more than one day.
The convexity of the light response curve of photosynthesis, was independent of whether the steady state measurements were made beginning in the dark and sequentially increasing photon flux density or beginning at high light and sequentially lowering photon flux density. Neither variation of CO₂ concentration from 35 to 200 Pa, nor of temperature from 5° to 32°C affected the convexity.     

The response of CO2 exchange rate to photosynthetic photon flux density for several Populus clones under laboratory conditions

No significant differences were found between four mathematical equations describing the response of CO₂ exchange rate to photosynthetic photon flux density in seven poplar clones under laboratory conditions. Choice of an optimal equation for poplar may be based on the contemplated aims. High significant differences (at p<0.001) were found among the clones.Research was supported by the Instituut tot Aanmoediging van het Wetenschappelijk Onderzoek in Nijverheid en Landbouw (I.W.O.N.L.), Brussels.     

Seasonal variation in hydrochloric acid, malic acid, and calciumions secreted by the trichomes of chickpea (Cicer arietinum)

A novel experimental set-up and method of recording of electrical potential differences in plants have been developed which enable continuous, 8-channel monitoring of electrical activity over extended periods of time using inserted, extracellular electrodes. The investigations were carried out on 21- to 23-day-old Helianthus annuus plants, and spontaneously-generated action potentials were recorded during monitoring sessions lasting for 3 days and nights. Characteristics of these spontaneous action potentials were elaborated, adopting as parameters their typical form, amplitude, duration, velocity, direction, and distance of propagation and frequency of occurrence in morphologically different parts of the plant, Variability, similarities, and interdependence of the above parameters in individual plants and in a group of 15 plants were determined. A hypothesis concerning propagation of action potentials in plants along specific impulse-propagating 'columns' is discussed. The frequency of generated impulses is highest at night and lowest in the day and also displays an apparent 24-h rhythm. Presumably this mechanism is under both endogenous and exogenous control and may be partly dependent on a biological clock.     

Role of roots and shoots in the regulation of the Fe efficiency responses in sunflower and cucumber

Several experimental approaches were used to study the role of roots and shoots in the regulation of the Fe efficiency responses in the roots of Fe deficient plants of sunflower ( Helianthus annuus L., inbred line RHA 274) and cucumber ( Cucumis sativus L., cv. Burpee pickler). The presence of the shoot apex and the youngest leaves in sunflower and cucumber plants was not indispensable to the development of the responses. In split-root experiments with sunflower plants, with either one or two shoots obtained by grafting, different parts of the root system could receive different Fe additions. Roots growing in a medium without Fe developed Fe efficiency re-sponses even when the other part of the root system was growing with Fe. Also, the part of the root system that was growing in a medium with Fe, developed some Fe efficiency responses, although delayed and at a lower level as compared with the root without external Fe. The results are discussed in relation to the idea that the development of the responses in the roots is controlled not only by the Fe content in the root, but also by some information transmitted between parts of a root system that differ in their Fe nutritional status.     

A comparison of nitrogen and carbon metabolism in the shallow and deep-water phytoplankton populations of a subalpine lake: response to photosynthetic photon flux density

Rates of , and CO₂ assimilationby the organisms in the shallow and deepchlorophyll layers ofCastle Lake were measured over a gradient of photosyntheticphoton flux densities (PPFD) during the 1979–1980 ice-freeseasons. The results of these experiments could be fitted witha hyperbolic function in the manner of the Michaelis-Mentenequation (excluding rates of dark assimilation) up to –40%of the surface PPFD after which photon inhibition occurred.The half saturation constants relative to incidence PPFD (K_LT)for assimilation ( = 1.1 E m^–2d^–2) were about twice those for ( = 0.5 E m^–2d^–1).All of the K_LT values correspond to depths in thelakerangingfrom 17–29 m(–1% of surface PPFD). Dark assimilationof both and was –50% of the assimilation at saturating PPFD implying that part ofthe immediate energy required for inorganic nitrogen assimilationmay come from intermediary metabolism. This contention was supportedfor assimilation by the results of experiments performed with specific inhibitors of non-cyclic photophosphorylationand oxidative phosphorylation. The K_LT values for the assimilationof CO₂ were from 2–10 times higher than those for inorganicnitrogen. These values for CO₂ assimilation were not significantlyaltered by the addition of either or during 12 h incubations.     

The dominant Basilicum Leaf mutation of sunflower controls leaf development multifariously and modifies the photosynthetic traits

Despite the remarkable morphological diversity in the leaf shape, usually higher plants maintain a flat lamina to ensure the desirable surface-to-volume ratio. The control of expansion of a flattened lateral organ is only partially known and the characterization of leaf mutants can be a useful strategy to add new information, as demonstrated in dicots with recessive mutants of model species. In sunflower, the dominant mutant Basilicum Leaf (BL) is characterized by prominent downward leaf curling. The central end of the present work is the detailed study of the BL phenotype to characterize the effects of this mutation, at organ and cellular levels, on both morphological and physiological traits. The geometry of mature leaves is obviously altered in BL plants because the lamina surfaces are downward curled and display a different length/width ratio with respect to wild type. The area of BL leaves is higher than that of wild type while the BL leaf perimeter is reduced. In BL leaves, the perimeter to the square root of area ratio is also abnormally low compared to the value of a flat sunflower lamina. These results suggest that the dominant mutation induces a deep change of the ratio between lamina and marginal growth. In addition, the epidermis and mesophyll of BL leaves comprised higher size pavement and palisade cells compared to those of wild type, indicating that the BL gene also control cell shape changes in the formation of the leaf lamina. Other leaf traits such as photosynthetic pigment content, pattern of minor vein architecture, margin type and stomatal density are affected by the mutation. In BL plants, the longitudinal axis of both cotyledons and ray flower corollas shows an unusual curvature while the diameter of shoot and inflorescence meristems is reduced. In mutant leaves, the gas exchange performance is influenced by low stomata conductance while the CO₂ assimilation rate is also depressed by a depleted Rubisco activity. Together, these data demonstrate the pleiotropic nature of the BL mutation, which affects a number of morphological aspects of flattened lateral organs as well as physiological parameters in sunflower.     

The effects of photoinhibition on the photosynthetic light-response curve of green plant cells (Chlamydomonas reinhardtii)

The photosynthetic response to light can be accurately defined in terms of (1) the initial slope (quantum yield); (2) the asymptote (light-saturated rate); (3) the convexity (rate of bending); and (4) the intercept (dark respiration). The effects of photoinhibition [which damages the reaction centre of photosystem II (PSII)] on these four parameters were measured in optically thin cultures of green plant cells (Chlamydomonas reinhardtii). The convexity of the light-response curve decreased steadily from a value of 0.98 (indicating a sharply bending response) to zero (indicating Michaelis-Menten kinetics) in response to increasing photoinhibition. Photoinhibition was quantified from the quantum yield of inhibited cells relative to that of control cells. The quantum yield was estimated by applying linear regression to low-light data or by fitting a non-rectangular hyperbola. Assuming the initial slope is linear allowed comparison with earlier work. However, as the convexity was lowered this assumption resulted in a significant underestimate of the true quantum yield. Thus, the apparent level of photoinhibition required for a zero convexity and the initial decrease in light-saturated photosynthesis depended upon how the quantum yield was estimated. If the initial slope of the light response was assumed to be linear the critical level of inhibition was 60%. If the linear assumption was not made, the critical level was 40%. At the level of inhibition where the convexity reached zero, the light-saturated rate of photosynthesis also began to decrease, indicating that this level of inhibition caused photosynthesis to be limited at all light intensities by the rate of PSII electron transport. At this level of inhibition the F_m-F_i signal (where F_m is maximal chlorophyll fluorescence and F_i is intermediate chlorophyll fluorescence of dark adapted cells; Briantais et al. 1988) from the fluorescence induction curve was zero and the F_i-F_o signal (where F_o is initial chlorophyll fluorescence of dark adapted cells) was 30% of the control, indicating dramatic reduction or complete elimination of one type of PSII. These data do not contradict published mathematical models showing that the ratio of the maximum speed of electron transport in PSII relative to the maximum speed of plastoquinone electron transport can determine the convexity of the photosynthetic response to light.Abbreviations and Symbols Chl chlorophyll content - DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea - F_o, F_i, F_m initial, intermediate, and maximal Chl fluorescence of dark adapted cells - P rate of net photosynthesis per unit chlorophyll (mol-(mg Chl)^–1 · s^–1) - PSII photosystem II - PQ plastoquinone - initial slope to the light-response curve - convexity (rate of bending) of the light-response curve of photosynthesis - Q photosynthetically active photon flux density (400–700 nm, mol · m^–2 · ^–1) The present investigation was supported by the Swedish Council for Forestry and Agricultural Research, the Swedish Environmental Protection Board, and the Swedish Natural Science Research Council. We thank Dr. Deborah D. Kaska (Department of Biological Sciences, University of California, Santa Barbara, Calif., USA) for giving us Chlamydomonas algae. We thank Professor G. Öquist (Department of Plant Physiology, University of Umea, Umea, Sweden) for his encouragement, valuable comments and discussion.