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.     

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).     

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.     

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.     

Modelling stomatal conductanceof field-grown sunflower under varying soil water content and leafenvironment: comparison of three models of stomatal response toleaf environment and coupling with an abscisic acid-based modelof stomatal response to soil drying

Stomatal conductance, g_s, responds both tothe immediate or local environment of the leaf, such as CO₂ partialpressure and irradiance, and to root‐sourced signals of water stress,particularly abscisic acid (ABA). Two models for the combined controlof g_s were formulated and tested in sunflower(Helianthus annuus). First, several empirical models weretested for the local control, demonstrating that the Ball–Berrymodel [Ball, Woodrow & Berry (in Progress in PhotosynthesisResearch Vol. 4, pp. 5.221–5.224: M. Nijhoff,Dordrecht, The Netherlands) 1987] is consistently amongthe most accurate. A problem of statistical non‐independence inthis model is shown to be minor. The model offers regularity ofparameter values among most species and, despite an oversimplicationin representing known humidity‐response mechanisms, it incorporates othersignalling loops from CO₂ and assimilation. In the firstcombined model, ABA as its concentration in xylem sap, [ABA]_xy,down‐regulates the slope, m, in the Ball–Berry modelby the factor g_fac = exp(– β[ABA]_xy).The ABA‐induced reduction in g_s decreases CO₂ assimilation andsurface humidity, thus appearing to induce the local‐control mechanismto amplify the ABA‐induced stomatal closure. In the second combinedmodel, g_s is estimated as the minimum of the local(Ball–Berry) response and the product g_fac g_s,max,with g_s,max as a maximal unstressed conductance.Both models can predict g_s from the external environmentalvariables with good accuracy (r² near 0·8 over20‐fold variations in g_s). Further analyses showthat g_s responds to humidity almost quadraticallyrather than linearly. It also responds to assimilation as a powerlaw with an exponent that is significantly less than 1. These limitations,shared by other models, suggest more research into biochemical signalling.     

During photosynthetic induction,biochemical and stomatal limitations differ between Brassica crops

Interventions to increase crop radiation use efficiency rely on understanding of how biochemical and stomatal limitations affect photosynthesis. When leaves transition from shade to high light, slow increases in maximum Rubisco carboxylation rate and stomatal conductance limit net CO₂ assimilation for several minutes. However, as stomata open intercellular [CO₂] increases, so electron transport rate could also become limiting. Photosynthetic limitations were evaluated in three important Brassica crops: Brassica rapa, Brassica oleracea and Brassica napus. Measurements of induction after a period of shade showed that net CO₂ assimilation by B. rapa and B. napus saturated by 10 min. A new method of analyzing limitations to induction by varying intercellular [CO₂] showed this was due to co-limitation by Rubisco and electron transport. By contrast, in B. oleracea persistent Rubisco limitation meant that CO₂ assimilation was still recovering 15 min after induction. Correspondingly, B. oleracea had the lowest Rubisco total activity. The methodology developed, and its application here, shows a means to identify the basis of variation in photosynthetic efficiency in fluctuating light, which could be exploited in breeding and bioengineering to improve crop productivity.     

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.     

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     

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.     

In vivo and in vitro effects of boron on the plasma membrane proton pump of sunflower roots

The effect of boron excess and deficiency on H⁺ efflux from excised roots from sunflower ( Heliarahus annuus L. cv. Enano) seedlings and on plasma membrane H⁺-ATPase (EC 3.6.1.35) in isolated KI-washed microsomes has been investigated. When seedlings were grown in media with toxic levels of H₃BO₃ (5 m M ) or without added boron and exposed to light conditions, an inhibition of the capacity for external acidification by excised roots was observed as compared to roots from seedlings grown with optimal H₃BO₃ concentration (0.25 m M ). Toxic and deficient boron conditions also inhibited the vanadate-sensitive H⁺-ATPase of microsomes isolated from the roots. The mechanism of boron toxicity was investigated in vitro with microsorne vesicles. A strong effect of boron on the vanadate-sensitive, ATP-dependent H⁺ transport was found, but the vanadate-sensitive phospho-bydrolase activity was not affected. These results suggest that boron could exert an effect on the plasma membrane properties, directly or indirectly regulating, proton transport.     

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.     

Analysis of the relationship between photosynthetic photon flux density and natural Taxus baccata seedlings occurrence

The aim of the present work was to analyse the relationship of seedlings and saplings of Taxus baccata to the photosynthetic photon flux density (PPFD) reaching the forest floor under natural conditions. Two permanent plots, subdivided into 1 × 1 m square plots, were established in a naturally regenerating population of T. baccata formed during last decades in the Kórnik Arboretum, Poland. All seedlings in every 1 × 1 m plots were counted. Relative PPFD was measured for every plot at the canopy height of the yew seedlings. The dependence of seedling density upon PPFD was examined. We found, that the frequency of the smallest seedlings (to 6.0 cm tall) was highest in the most shaded plots and decreased in plots with increasing PPFD. Thus, the youngest yew seedlings can germinate and grow in very shady conditions. However, the older seedlings (6.1–25.0 and 25.1–100.0 classes) were observed most frequently in 2–7% PPFD. The small numbers of older, taller seedlings in deep shade likely indicate a higher mortality rate of seedlings less than 6 cm in height without promotion to the next height class. Probably the low value of PPFD under the canopy of the stand significantly reduces the competition of other plants with the youngest yew seedlings. At higher light levels they may not be able to compete with more light-demanding plants, such as herbs and seedlings of broad-leaved trees. The seedlings of the second (6.1–25.0 cm) and third (25.1–100.0 cm) height classes were observed most frequently in the plots with 2–7% PPFD (Fig. 1b and c).     

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.     

四种荒漠植物的光合响应

研究了胡杨（Populus euphratica Oliv）、疏叶骆驼刺（Alhagi sparsifolia B.Keller et Shap)、多枝柽柳（Tamarix ramosissima Lbd)和头状沙拐枣（Calligonum caput-medusae Schrenk)等四种塔克拉玛干荒漠植物的光合响应曲线,结果表明：（1）四种植物中,疏叶骆驼刺光呼吸（Rp)和CO2补偿点（Г）最高,净光合速率（An）以及水分利用效率（WUE）最低;而C4植物头状沙拐枣无论Rp还是Г都最低,它的An、光补偿点（LCP）及光饱和点（LSP）都显著高于其它3种植物（P≤0．05）,而且因为非常低的蒸腾速率（E）,沙拐枣的WUE也较高。（2）4种植物都为喜光植物,但仍然表现出一定的差异。根据它们光曲线中LCP以及暗呼吸速率（Rd),4种喜光植物的喜光顺序为：头状沙拐枣,疏叶骆驼枣刺,多枝柽柳,胡杨。（3）柽柳光合响应曲线与日变化中的An,gs等值差异较大,可能是该植物的温度生态位较其它三种植物范围较窄,设置的25℃不在其最适温度范围内,或者是在这个温度下气孔对光强变化的不敏感。（4）综合前人的结果表明,在特定的环境条件,发育阶段以及经过一定的诱导处理,胡杨可以因诱导而表现出一些C4植物特征。     

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.     

Evaluating the usefulness of hemispherical photographs as a means to estimate photosynthetic photon flux density during a growing season in the understorey of Nothofagus pumilio forests

Abstract

Old-growth Nothofagus pumilio forests in Chile are managed employing a shelterwood system. A wide range of canopy openings can be found in old-growth and managed forests. Plant survival and growth in the understorey are influenced by the light available. There are limitations (practical and economic) to monitoring the light in the understorey. The aim of this study was to assess the options to estimate the forest understorey photosynthetic photon flux density (PPFD) measured during the growing season (GS) using canopy openness (CO) estimated by means of hemispherical photographs (HP). PPFD was measured using 31 sensors (Li-190SA quantum sensor) over the course of three GSs (October to March). The sensors were installed in an old-growth stand and another subjected to a regeneration felling under a shelterwood system. One HP was taken above each sensor (during the final GS) and the CO estimated. A comparison of the three seasons revealed that the sum of the PPFD during the GSs did not differ significantly. The CO could be used to effectively predict the sum of the PPFD during a GS (R ² = 0.959). These results demonstrate the usefulness of HPs as a means to estimate the sum of the PPFD during a GS.