Antagonistic but complementary actions of phytochromes A and B allow seedling de-etiolation.

Using dichromatic radiation, we show that the actions of phytochromes A and B (phyA and phyB) in Arabidopsis thaliana are antagonistic in mediating red and far-red radiation effects on seedling de-etiolation and yet act in a complementary manner to regulate de-etiolation, irrespective of spectral composition. At low phytochrome photoequilibria inhibition of hypocotyl extension was strong, because of the action of a far-red high-irradiance response mediated by phyA. At high phytochrome photoequilibria inhibition of hypocotyl extension was also strong, because of the action of phyB. At intermediate photoequilibria hypocotyl inhibition was less strong. In their natural environment, this dual action will strongly retard hypocotyl growth and promote cotyledon opening and expansion both in open daylight and under dense vegetation. Overlapping action by phyA and phyB will substantially promote de-etiolation in sparse vegetation. The antagonistic and complementary actions of phyA and phyB, therefore, allow the optimum regulation of seedling growth after emergence from the soil.     

Phytochrome Control of Chlorophyll Content in Mature Attached Leaves of Petunia axillaris

Petunia axillaris plants were grown under white light (photoperiod10 h). Pulses of end-of-day far-red light, but not red lightat the end of the day slowed down the accumulation of chlorophyllin expanding, greening leaves and reduced chlorophyll contentin mature leaves. The chlorophyll a / b ratio was unaffected.Low phytochrome photoequilibria reduced chlorophyll contentin mature leaves without affecting leaf area or d. wt. Thiseffect occurred even before senescence was triggered in thecontrol plants having high phytochrome photoequilibria     

Photomorphogenically Defined Light and Resistance of Poa pratensis to Drechslera sorokiniana

Photomorphogenic light definitions were derived by mathematical determination of the estimated phytochrome photoequilibrium for each light treatment spectrum. A wide range of photomorphogenic light treatments represented by spectra with estimated phytochrome photoequilibria of 0.45, 0.54, 0.60, 0.67, and 0.71 was utilized to determine the influence of photomorphogenically defined light on resistance of Poa pratensis L. to pathogenesis by Drechslera sorokiniana. Accurate resolution of D. sorokiniana leaf spot development required evaluation of separate leaf ages due to the sequential appearance, development, and senescence of P. pratensis leaves. Disease development (all light treatments) was greatest on leaf 4 (oldest, postmature) followed by leaf 1 (youngest, premature). Low levels of disease occurred on leaves 2 and 3 (mature). Photomorphogenic light defined by estimated phytochrome photoequilibria greater than 0.60 (natural light 0.60) was most disease promotive on leaf 1. Conversely, photomorphogenic light defined by estimated phytochrome photoequilibria of less than 0.60 was most disease promotive on leaf 4. These responses indicate that inherent resistance or susceptibility expressed by P. pratensis to pathogenesis by D. sorokiniana is regulated in part by leaf age (developmental senescent stage) and by photomorphogenically defined light quality. A hypothesis is presented and discussed which integrates and speculates on these observations with respect to the literature.     

On phytochrome absorption and the phytochrome photoequilibrium in a green leaf: environmental sensitivity and photoequilibrium time

The average, corrected attenuance spectra for both spectral forms of phytochrome in a mature leaf were calculated. Optical masking by chlorophyll together with the detour effect (optical path lengthening effect) due to multiple light scattering led to large changes in both the Qy band shape and wavelength position and the effective intensity of the weak vibrational bands increases. The Pfr/Pr oscillator-strength-ratio between 400-750 nm (0.93 in vitro), becomes 1.63 in a leaf. Thus the dominant absorption form is Pfr. These two values permit calculation of the phytochrome photoequilibrium under conditions of "daylight" illumination both in vitro and in folia. These values are 0.6 and 0.38 respectively. Previous literature estimates for the situation in vitro, based on the 660/730 nm absorption ratio, yielded values close to 0.6. It is demonstrated that this large decrease in the phytochrome photoequilibrium in a leaf has the effect of translating this parameter to a position on the dose (red/far-red light ratio)-response (Pfr/Ptot) plot towards greater sensitivity to changes in the environmental red/far-red ratio. The increased sensitivity factor is almost five-fold for the "daylight" environment and is even greater for the various "shade-light" environments. The approximate time taken to attain photoequilibrium (1/e lifetime) has also been calculated for phytochrome in a leaf in different light environments. For the "daylight" environment the photoequilibration time is approximately 5 s, which increases into the 20-80 s interval under different degrees of "shade light". Thus, despite the strong optical masking by chlorophyll in a mature leaf, the phytochrome photoequilibrium is attained quite rapidly on a physiological time scale.     

COMPARATIVE FLOWER DEVELOPMENT IN THE CLEISTOGAMOUS SPECIES VIOLA ODORATA. I. A GROWTH RATE STUDY

In Viola odorata, chasmogamous (CH) or open flowers and small, short-petioled leaves are produced under 11 hr or less of daylight, cleistogamous (CL) or closed flowers and large, long-petioled leaves under 14 hr or more of daylight, and intermediate floral and leaf forms under transitional photoperiods. CL flowers are approximately four times smaller than CH flowers and differ morphologically in repressed growth of the anterior petal spur and staminal nectaries, and recurving of the style which remains enclosed within the cone formed by anther appendages. Both CH and CL shoot systems conform to a (2 + 3) phyllotaxis with minor differences in leaf divergence angles and phyllotactic indices. The larger CL leaf grows significantly faster than the CH leaf, and an increased rate of leaf initiation occurs in the CL apex represented by a plastochron of 3.4 days compared to 4.3 days in the CH apex. The plastochron index was used to indirectly age young floral primordia nondestructively until prophase of meiosis I within the anthers. This event occurs 8 days earlier in the CL than the CH flower. Time from meiosis until flower maturity, determined by direct observation, is about 14 days for the CL flower, versus 21 days for the CH flower.     

绒毛番龙眼对生长光强的形态和生理适应

在100％、50％、25％和8％自然光强下栽培绒毛番龙眼幼苗并研究了其对光环境的适应。100％生长光强下绒毛番龙眼通过增大叶片悬挂角(midrib angle,MA)和比叶重(lamina mass per unit area,LMA),减少叶氮在捕光组分中的分配等降低光能捕获;通过增加类胡萝卜素含量增加热耗散。虽如此,还是发生了比较严重的光抑制,加之叶氮在光合机构中的分配最少,导致光合能力最低,长势最差。8％生长光强下绒毛番龙眼通过降低MA、LMA以及叶片技转,增加叶氮在捕光组分中的分配等提高光能捕获能力,光能转换及利用效率较高,热耗散水平较低,但由于环境光较弱,限制了光合碳同化,植株生长也较慢。50％和25％生长光强下绒毛番龙眼有较强的光能捕获、利用和耗散能力,在几种光处理中长势最好。     

Phytochrome control of the growth and development of Rumex obtusifolius under simulated canopy light environments*

Abstract Studies on the growth and development of Rumex obtusifolius seedlings under simulated shade conditions show that both light quantity and quality contribute to the observed responses. In the shade situation the plants have lower dry matter, lower leaf area and lower net assimilation rates. Petiole elongation occurs under shade conditions only after transfer of the plants from non-shade environments. The effects of light quality are related to phytochrome photoequilibria set up by the increased relative photon flux in the far-red found under vegetation canopies.     

The induction of sun and shade leaves of the European beech (Fagus sylvatica L.): anatomical studies

Summary Primordia from buds of sun and shade twigs of European beech (Fagus sylvatica L.) were collected six times a year for anatomical investigations. Differentiation into sun-leaf and shade-leaf primordia was first observed in early August. Sun-leaf primordia had five, and shade-leaf primordia four layers of mesophyll meristem cells. With potted graft unions of beeches possible structural changes of leaf primordia were investigated. Trees adapted to shade develop sun-leaf primordia when put into full daylight, provided the transfer happened before July. Trees adapted to full daylight developed leaf primordia which remained structurally sun-leaf primordia when the plant was kept under shade conditions. Shadeleaf branches of young beech trees cut in February in order to expose the shade buds to full daylight developed either shade leaves or intermediate shade/sun leaves. These experiments show that the subtending leaf may provide the developing axillary bud with photoassimilates, but its character, whether sun or shade leaf, has no influence on the character of the developing leaf primordia.     

A simple method to estimate photosynthetic radiation use efficiency of canopies

BACKGROUND AND AIMS: Photosynthetic radiation use efficiency (PhRUE) over the course of a day has been shown to be constant for leaves throughout a general canopy where nitrogen content (and thus photosynthetic properties) of leaves is distributed in relation to the light gradient. It has been suggested that this daily PhRUE can be calculated simply from the photosynthetic properties of a leaf at the top of the canopy and from the PAR incident on the canopy, which can be obtained from weather-station data. The objective of this study was to investigate whether this simple method allows estimation of PhRUE of different crops and with different daily incident PAR, and also during the growing season. METHODS: The PhRUE calculated with this simple method was compared with that calculated with a more detailed model, for different days in May, June and July in California, on almond (Prunus dulcis) and walnut (Juglans regia) trees. Daily net photosynthesis of 50 individual leaves was calculated as the daylight integral of the instantaneous photosynthesis. The latter was estimated for each leaf from its photosynthetic response to PAR and from the PAR incident on the leaf during the day. KEY RESULTS: Daily photosynthesis of individual leaves of both species was linearly related to the daily PAR incident on the leaves (which implies constant PhRUE throughout the canopy), but the slope (i.e. the PhRUE) differed between the species, over the growing season due to changes in photosynthetic properties of the leaves, and with differences in daily incident PAR. When PhRUE was estimated from the photosynthetic light response curve of a leaf at the top of the canopy and from the incident radiation above the canopy, obtained from weather-station data, the values were within 5 % of those calculated with the more detailed model, except in five out of 34 cases. CONCLUSIONS: The simple method of estimating PhRUE is valuable as it simplifies calculation of canopy photosynthesis to a multiplication between the PAR intercepted by the canopy, which can be obtained with remote sensing, and the PhRUE calculated from incident PAR, obtained from standard weather-station data, and from the photosynthetic properties of leaves at the top of the canopy. The latter properties are the sole crop parameters needed. While being simple, this method describes the differences in PhRUE related to crop, season, nutrient status and daily incident PAR.     

Vegetative growth and leaf resin composition in Hymenaea courbaril under photoperiodic extremes

Seedlings from five geographic populations of the widespread Hymenaea courbaril were grown for one year under two extremes of photoperiod. The short photoperiod treatment consisted of 8 hr of unsupplemented greenhouse daylight and the long photoperiod treatment of greenhouse daylight at the natural photoperiod of Santa Cruz, California (37° N) plus a 3-hr night interruption (0.25 mW/cm²). Growth was measured by total stem length and leaf resin sesquiterpene composition was quantitatively analyzed by GLC. Photoperiod had no significant effect on leaf resin chemistry but the longer photoperiod markedly promoted vegetative growth. Large population differences in overall vegetative growth and quantitative leaf resin composition were also evident. Growth patterns indicated that the low intensity night break had little or no effect on growth.     

Growth and Development of the Banana Plant Gross Leaf Emergence

The banana leaf must pass through a prolonged growth periodwhile totally within the pseudostem and with the lamina in aninvolved double coil. During this time the blade makes mostof its growth, completing the last fourth of the expansion asit emerges. In this same latter period it accomplishes almosttwo-thirds of its elongating growth. The first few leaves of a single plant are essentially bladeless.Thereafter the size of the lamina increases in both dimensionswith each succeeding leaf tending to exceed its predecessor.A diurnal/nocturnal cycle is evident in both Costa Rica andHonduras and maximal elongation coincides with the highest temperaturein the daylight hours. Control of elongation is elegant witheach leaf developing at a rate governed by those ahead of it.An assessment is made of the stomatal occurrence over the surfaceof the blade.     

毛白杜鹃‘紫萼’的光适应性及最适光强的研究

从形态、生理角度研究了杭州园林中应用最广泛的杜鹃‘紫萼’(Rhododendron mucronatum cv Plenum)的光适应性和最适光强生境。结果表明:随着叶片遮荫程度的增加,杜鹃的叶面积和叶绿素含量增加;光补偿点、光饱和点及暗呼吸强度下降,说明杜鹃对弱光生境有一定的适应性。另一方面,随着相对光强的增加,叶片厚度,比叶重以及栅栏组织、海绵组织厚度及其比值,可溶性蛋白质及净光合速率增加,表现出对阳生生境更好的适应性。在生境65％全光照时,植株在形态,解剖及生理上均处于最佳状态。因此,65％全光照的生境是毛鹃‘紫萼’的最佳光生境。     

Variations in the shallow water form of Potamogeton richardsonii induced by some environmental factors

SUMMARY. Populations of Potamogeton richardsonii in Sparrow Lake, Ontario, vary greatly in leaf dimensions and internode length. Leaf length/breadth (L/B) ratio is increased by low irradiance, significantly at 4% daylight and, in a contrary fashion, by ontogenetic drift at high irradiances of shallow water. L/B ratio was not correlated with substrate. These findings suggest that this ratio is of doubtful value taxonomically or as an integrator, as specific leaf area is confirmed to be, of factors like photosynthetically active radiation (PAR). Longest internodes belong to plants from deep(1.8 m) water, and, in a summer-grown high-density population, internode length decreases logarithmically as depth lessens; both observations implicate PAR and ageing. Relative to undisturbed, shallow-water shoots, young transplants in full daylight elongate almost twice as fast because of the production of more, and longer, internodes. Young shoots in 12% daylight lengthen even more rapidly than those in full daylight (and four times faster than undisturbed shoots) in the same period because of quicker elongation of the same number of internodes. At ambient summer temperatures, this rate of elongation is inversely related to PAR. Experimental and seasonal field data indicate that maximal internode extension occurs on young shoots in very low irradiances at temperatures of 9–15°C; in nature, effects of increasing water temperatures are depressed by increasing irradiances and ageing until minimal extension takes place in old shoots at mid- to late-summer temperatures and in high irradiances. In tanks (40 m³) in full daylight, nutrients limit growth of plants on sand before they limit growth on clay (with marl intermediate) whereas, at 12% daylight, light limits growth on sand before soil nutrients do. The leaf area index on silty sand on an exposed shore was 0.4 at 0.5 m depth, in contrast to a cultured population where it was 4.0 in silty loam at the same depth. Thus, while it has not been possible to explain some variations in leaf morphology of P. richardsonii in environmental terms, the differential effects of ontogeny, PAR and temperature on shoot growth have been assessed, along with overall effects of light and nutrients on biomass and of shelter on leaf area index.     

Adaptability ofMentha piperita l. to irradiance. Growth,specific leaf area and levels of chlorophyll,protein and mineral nutrients as affected by shading

Height growth ofMentha piperita L. is better at 44% than 100% daylight. At 14% daylight, growth is limited, probably by lack of photosynthate, and no flowering occurs. Specific leaf area and protein content increase significantly with the decrease of irradiance. Shade leaves contain more chlorophyll and potassium than sun leaves. Sun leaves show higher levels of calcium because of their more xeromorphic structure. Transfer experiments between light and shade indicate thatM. piperita responds very effectively to irradiance also in a late stage of the life cycle with changes of specific leaf area and chlorophyll content.     

Apoptosis in Wheat Seedlings Grown under Normal Daylight

Apoptosis was observed in the coleoptile and initial leaf in 5-8-day-old wheat seedlings grown under normal daylight. Apoptosis is an obligatory event in early wheat plant ontogenesis, and it is characterized by cytoplasmic structural reorganization and fragmentation, in particular, with the appearance in vacuoles of specific vesicles containing intact organelles, chromatin condensation and margination in the nucleus, and internucleosomal fragmentation of nuclear DNA. The earliest signs of programmed cell death (PCD) were observed in the cytoplasm, but the elements of apoptotic degradation in the nucleus appeared later. Nuclear DNA fragmentation was detected after chromatin condensation and the appearance in vacuoles of specific vesicles containing mitochondria. Two PCD varieties were observed in the initial leaf of 5-day-old seedlings grown under normal daylight: a proper apoptosis and vacuolar collapse. On the contrary, PCD in coleoptiles under various growing (light) conditions and in the initial leaf of etiolated seedlings is only a classical plant apoptosis. Therefore, various tissue-specific and light-dependent PCD forms do exist in plants. Amounts of O2*- and H2O2 evolved by seedlings grown under normal daylight are less than that evolved by etiolated seedlings. The amount of H2O2 formed in the presence of sodium salicylate or azide by seedlings grown under normal daylight was increased. Contrary to etiolated seedlings, the antioxidant BHT (ionol) did not inhibit O2*- formation and apoptosis and it had no influence on ontogenesis in the seedlings grown under normal daylight. Thus, in plants grown under the normal light regime the powerful system controlling the balance between formation and inactivation of reactive oxygen species (ROS) does exist and it effectively functions. This system is responsible for maintenance of cell homeostasis, and it regulates the crucial ROS level controlling plant growth and development. In etiolated plants, this system seems to be absent, or it is much less effective.     

Adaptations of the Photosynthetic Mechanism of Sugar Maple (Acer saccharum) Seedlings Grown in Various Light Intensities

Sugar maple (Acer saccharum Marsh.) seedlings were grown in a nursery for three years in 13, 25, 45 and 100 per cent of full daylight. During the third year of growth, the rates of their apparent photosynthesis and respiration were measured periodically with an infra-red gas analyzer at various light intensities and normal CO₂ concentration. In addition, the rates of apparent photosynthesis of a single attached leaf of the same seedlings were measured at saturating light intensity, hut varying CO₂ concentrations. An increase in the light intensity in which seedlings were grown had no effect on their height or mean leaf area, hut resulted in thicker leaves, an increase in the total leaf area per seedling due to an increase in the number of leaves, an increase in the dry weight especially of roots and a decrease in the chlorophyll content of leaves. Throughout the growing season seedlings grown in full daylight, as compared with those grown in lower light intensities, had the lowest rates of apparent photosynthesis measured at standard conditions (21,600 lux light intensity and 300 ul/l of CO₂), when this was expressed per unit leaf area, hut the highest rates on a per seedling basis. Thus dry matter production attained at the end of the growing season correlated positively with the photosynthetic rate per seedling, but not per unit leaf area. The rates of apparent photosynthesis of seedlings grown at lower light intensities were more responsive to changes in light intensity or CO₂ concentration than those of seedlings grown in full daylight intensity.     

Some observations on the spectral distribution characteristics of short-wave radiation within Pinus radiata D. Don canopies

Abstract. Measurements of the photosynthetic photon flux density (400–700 nm) and of the spectral distribution of photon flux density across the 370–800 nm waveband, were made under both clear and overcast sky conditions above and at various positions within two Pinus radiata canopies of different stocking but similar leaf area indices. The spectra obtained for the daylight conditions (i.e. above forest canopy) were generally similar to those published previously. The spectra for shadelight within the forest canopy showed no blue peak which was characteristic of previously reported measurements which were restricted to the diffuse radiation component. There was almost neutral absorption within the 400–700 nm waveband, and typical lower attenuation in the 700–800 nm waveband. The blue: red ratio was largely unchanged by either canopy type or sky conditions and varied between 0.57 and 0.81. The red: far-red ratio in shadelight was between 0.22 and 0.41 under clear sky and between 0.68 and 0.95 under overcast sky conditions. Values for daylight were between 1.16–1.22. Calculated phytochrome photoequilibrium values in shadelight were approximately 0.35 under clear sky and 0.46 under overcast sky conditions. In each case there appeared to be no differences between the two canopies with respect to these minimum values.     

A new life table of leaves ofPhaseolus vulgris L. in relation to their position within the canopy and plant density

In order to demonstrate in detail the relationship between the longevity and productivity of leaves within a canopy, a new life table approach, the ‘bioeconomic life table’, was applied to the leaves of kidney bean plants (Phaseolus vulgaris L.) in relation to planting density and their position within the canopy. The net photosynthetic rate for upper leaves under full daylight tended to decline gradually due to leaf senescence from about 20 days after leaf emergence, and for the lower leaves the decrease was very rapid due to both shading and senescence about 10 days after emergence. Analysis of the survivorship curves and daily surplus production of leaves suggested that the lower and middle leaves, especially the latter, survived without surplus production of dry matter after they had reached mean longevity, and while the upper leaves at high density had a much shorter mean longevity, they had very large values of daily surplus production throughout the survival period. For the total foliage, the summed value of accumulated surplus production during the survival period was about five times as large as the summed value of the dry weight of the dead leaves, regardless of planting density. The daily rate of canopy leaf respiration was almost proportional to that of canopy gross photosynthesis for the various leaf area indices of the canopy, so that there was no optimum leaf area index that maximized canopy daily surplus production.     

Regulation of photosynthate partitioning into starch in soybean leaves : response to natural daylight

Studies conducted in controlled environments indicate that daylength affects the proportion of photosynthate stored in leaves as starch or sucrose. To examine the response of partitioning to natural daylight, soybeans (Glycine max [L.] Merr. cv Williams) were grown at 12 different times between May and November in a constant temperature greenhouse without supplemental lighting. Plants were transferred from the greenhouse to a controlled environment chamber at the end of civil twilight at a set developmental stage (expanding seventh trifoliolate leaf, counting acropetally). Net photosynthesis and the accumulation of starch and sugar in fully expanded fourth trifoliolate leaves were determined the following day under standard conditions in the chamber (lights-on synchronized with sunrise). Photosynthesis on a leaf area basis decreased about 10% between midsummer and early autumn. Leaf soluble sugar accumulation was low at all harvests. However, a twofold increase in photosynthate partitioning into starch occurred over the same time period, resulting in an 80% increase in absolute starch accumulation rate. Starch was responsible for about 78% of the increase in leaf dry matter during the light at all harvests, indicating that starch accumulation as affected by prior daylight conditions will alter export of photosynthate during the light period. Photosynthate partitioning into starch was linearly correlated with daylength at harvest, prior average peak solar irradiance, and other parameters that correlated with daylength and solar radiation such as harvested top dry matter. The relation between growth and seasonal changes in daylight (including daylength, irradiance, and light integral) are discussed in relation to photosynthate partitioning under field conditions.     

Physiological and Ecological Studies in the Analysis of Plant Environment: IX. Adaptive Changes in the Vegetative Growth and Development of Helianthus annuus induced by an Alteration in Light Level

Pot experiments were carried out in which sunflowers in theearly vegetative phase were first grown for a period under threelevels of light (1·0, 0·5, and 0·24 daylight).Subsequently pots from each light group were subdivided intothree so that in a second period plants could be subjected tothe nine possible combinations of the same three light intensitiesbefore and after transference. During the post-transference period of adaptation to eithera higher or a lower intensity the net assimilation rate is logarithmicallyproportional to the light received and there is no residualeffect of the initial light treatments. Eight days after transferencethe leaf-area ratios (total leaf area/total plant weight) ateach light level become adjusted to a new equilibrium irrespectiveof the large initial differences in the ratio induced by thepre-transference intensities. In both periods there is an inverseand logarithmic relationship between the leaf-area ratio andfalling light intensity; consequently, the greater the degreeof shading in the pre-transference period, the higher are themean ratios in the second period. Since the relative growthrate is the product of the net assimilation rate and the leaf-arearatio, the variations in the leaf-area ratio in the post-transferenceperiod induced by the initial light treatments are reflectedin the relative growth rates. Thus plants transferred from alower to a higher light intensity are leafier and initiallygrow faster than plants maintained at the higher level in bothperiods: the converse conditions lead to a reduction in thegrowth rate. Shading depresses the growth of the roots, but the relativegrowth rate is dependent on the light intensity in both thepre-transference and post-transference periods. With transferencefrom daylight to 0·24 daylight, the roots during theperiod of adjustment may lose weight, while the growth rateis maximal when plants are moved from the lowest to the highestintensity. In two out of the three experiments the relativegrowth rate of the shoot in the post-transference period isof the same order at all light intensities and is largely independentof the light received in the initial period. In terms of leaf weight, decreasing the light intensity decreasesthe relative growth rate and there is no consistent after-effectof the initial light treatments. The rate of expansion in leafarea tends to be highest at the intermediate level of 0·5daylight and over all the post-transference intensities therates are maximal for those plants which received initiallyfull daylight. The ratio of leaf area to leaf weight is inversely and logarithmicallyproportional to the light level. After transference the slopesof the regressions are independent of the initial light treatments,but the mean ratios are inversely correlated with the initialdegree of shading. These adaptive changes to a variation in the light level arediscussed with particular reference to the control of growthexerted by growth-regulating substances. It is concluded thaton the basis of existing knowledge no adequate interpretationis yet possible.