Physiological and Ecological Studies in the Analysis of Plant Environment: VII. An Analysis of the Differential Effects of Light Intensity on the Net Assimilation Rate, Leaf-Area Ratio, and Relative Growth Rate of Different Species

Since relative growth rate is the product of net assimilationrate and leaf-area ratio (leaf area/plant weight), it followsthat if the effects of shading on both net assimilation rateand leaf-area ratio can be expressed mathematically, then therelationship between light intensity and relative growth ratecan be derived from the product of the two mathematical expressions. For all the ten species investigated in field and pot cultureexperiments, it has been found that during the early vegetativephase both the changes in leaf-area ratio and net assimilationrate, over the range of 0·1 to full daylight, are linearlyrelated to the logarithm of the light intensity. In consequence,the relationship between relative growth rate and the logarithmof light intensity—being the product of the two linearregressions—is curvilinear. For species of shady habitats (Geum urbanum, Solamun dulcamara)neither the levels of assimilation rate nor the ‘compensation-point’values are very different from those of the eight species fromopen situations (e.g. Hordeum vulgare, Pisum sativum, Fagopyrumesculentum). Nevertheless the intensity at which growth rateis maximal varies between species: it is 0•5 for G. urbanum,0•7 for H. annuus, full daylight for F. esculentum, whilefor Trifolium subterraneum the calculated value is 1·8daylight. Such specific differences can be largely accountedfor in terms of the differences in leaf-area ratio at the differentlight levels. On the basis of this analysis of the light factor, a ‘shade’plant is best redefined as a species in which a reduction ofthe light intensity causes a rapid rise in the leaf-area ratiofrom an initial low value in full daylight: for a ‘sun’plant the converse definition holds.     

Interrelationships between Light Intensity and the Physiological Effects of 2:4-Dichlorophenoxyacetic Acid on the Growth of Helianthus annuus

The interrelationships between light intensity and the effectsof sodium 2:4-dichlorophenoxyacetate on the growth of Helianthusannuus in the early vegetative stage have been studied by subjectingtreated and control plants to a range of light intensities from,1.0 to 0.12 daylight. The growth regulator in varying amountswas applied either as droplets of aqueous solution to the firstor second pairs of leaves or as an overall spray. When the amountof the compound is such as to cause small but significant reductionsin the relative growth rate, the leaf-area ratio and the ratioof leaf area to leaf weight are likewise depressed but the netassimilation rate is relatively unaffected. Between 0.25 daylightand full daylight the proportionate changes in the relativegrowth rate, leaf-area ratio, and net assimilation rate inducedby sub-lethal amounts of the growth regulator are not greatlymodified by the level of light. If the intensity is reducedfurther to 0.12 daylight, then the reactions of the shaded plantsdiffer markedly from those of unshaded plants. For example,when the plants are shaded both before and after a spray application,the concentration required to cause a 50 per cent. mortalityis one-tenth of that demanded for plants receiving full daylight.For less phytotoxic amounts the percentage reductions, relativeto the controls, induced in the growth rate of the shoot aremore dependent on the intensity after than before the applicationof either a spray or measured droplets. These greater depressions in shoot growth at 0.12 daylight afterthe application are linked with comparable depressions in therate of growth of the leaves of which the first pair are moresensitive. Similarly, the growth rate of the first internodeis also depressed more under shade conditions, but that of thesecond internode at low doses may be greatly increased. Forboth the leaves and internodes shading before as against shadingafter the application may have different significant effectson the changes in growth caused by the growth regulator. Undersome conditions the interactions between the light intensitybefore and after the application and the quantity of the growthregulator are exceedingly complex. By using sodium 2:4-dichloro-5-iodophenoxyacetatelabelled with radioactive I¹³¹, it has also been establishedthat the rate of penetration into leaves is accelerated by ahigh light intensity after the application, but that the intensityreceived prior to the application causes no significant effect.Evidence was also obtained that transport to the shoot fromthe treated leaf is more dependent on the light intensity afterthe application—the transport being greater in full daylight.It is concluded that a number of factors must be involved inbringing about these differential effects and their relativeimportance is discussed.     

Physiological and Ecological Studies in the Analysis of Plant Environment: X. An Analysis, of the Effects of Seasonal Variation in Daylight and Temperature on the Growth of Helianthus annuus in the Vegetative Phase

To determine on a quantitative and mathematical basis the effectsof seasonal changes in the levels of daylight and temperatureon vegetative growth and development in two years pot experimentsin the open were carried out at successive weekly intervalsbetween May and September. So as to minimize errors arisingfrom ontogenetic drifts the procedure adopted was to sow atintervals of a few days throughout the season batches of potswith seed of Helianthus annuus and to select pots containingplants of a standard morphological status for the start of eachweekly experiment. At the beginning and end of the week halfthe pots were harvested, the plants divided into root, stem,and leaf, the leaf area estimated, and the dry weights determined.The diurnal changes in air temperature were continuously recordedwhile the amount of daylight, excluding infra-red and ultravioletradiation, was measured with a specially constructed integratingrecorder. From the biological data for each week twelve variables werecalculated, namely the relative growth rates of both the wholeplant and the individual parts, the proportion by dry weightof the individual parts (root-, stem-, and leaf-weight ratios),the ratio of leaf area to total plant weight (leaf-area ratio),the rate of leaf expansion, the ratio of leaf area to leaf weight,and the net assimilation rate on the criteria of leaf area andweight. The main independent variables considered were the meanweekly temperature, the mean daily maximum minus the mean nightlyminimum temperature, the total amount of light per week, andthe time of year when the individual experiment was undertaken. Multiple regression analyses showed that (i) save for the stem-weightratio the data for the two years could be pooled, (ii) the fluctuationin diurnal temperature was of little account, (iii) transformationof the light data to either logarithms or square roots did notimprove the fit and (iv) for some of the dependent variables,e.g. leaf-area ratio, the ‘time of year’ effectwas significant but could be eliminated if the equation wasmodified to predict the value at the end of the week, giventhe initial value and the light and temperature data. The final series of multiple regressions revealed that (i) theleaf-weight ratio is not controlled by either the amount oflight or mean temperature, (ii) the relative growth rate ofthe root and the root-weight ratio are positively linked onlywith temperature, (iii) the rate of leaf growth either in areaor weight together with the net assimilation rate (area basis)are positively dependent on light alone, (iv) the net assimilationrate (weight basis) and the relative growth rates of the wholeplant and the stem are directly and positively correlated withboth temperature and light, and (v) the leaf-area ratio, theratio of leaf area to leaf weight and the stem-weight ratioare depressed by increasing light but augmented by rising temperature.In the individual regressions for net assimilation rate (areaand weight), the relative growth rates of the whole plant, stemand leaf weight, and the ratios of stem weight and leaf areato leaf weight the percentage variation accounted for rangedfrom 47 to as high as 91 per cent. The implication of these findings in relation to experimentsin controlled environmental chambers are discussed.     

Physiological and Ecological Studies in the Analysis of Plant Environment: VI. The Constancy for Different Species of a Logarithmic Relationship between Net Assimilation Rate and Light Intensity and its Ecological Significance

In this further study of light as an environmental factor theeffects of shading on the growth of Helianthus annuus, Fagopyrumesculentum, Trifolium subterraneum, Tropaeolum majus, Lycopersicumesculentum, Vicia faba, Pisum sativum, Hordeum vulgare, Solanumdulcamara, and Geum urbanum have been investigated. It has been established that the reductions in the net assimilationcaused by shading are similar for all ten species. The net assimilationrate during the season of active growth is linearly relatedto the logarithm of the light intensity. When similar experimentsare conducted late in the autumn and the relative growth rateis very low, the logarithmic relationship no longer holds. From the data it is possible to obtain precise estimates ofthe compensation point. The mean values of the compensationpoint ranged from 0·06 to 0·09 of daylight foreight species, while for V. faba and H. vulgare somewhat higherfigures were obtained—0·14 and 0·18 of daylight. Flctuations in the net assimilation rate in full daylight showedno correlation with variations in the value of the compensationpoint. From these results it is concluded that species cannot be groupedinto ‘sun’ or ‘shade’ plants, eitheron the basis of differences in the value of the compensationpoint or on the grounds that there are large variations in theeffects of shading on net assimilation rate.     

Physiological and Ecological Studies in the Analysis of Plant Environment: XIII. A Comparison of the Effects of Seasonal Variations in Light Energy and Temperature on the Growth of Helianthus annuus and Vicia faba in the Vegetative Phase

The influences of seasonal changes in light radiation and temperatureon the vegetative growth of Helianthus annuus and Vicia fabahave been investigated in the east of Scotland by pot experiments,carried out in the open at weekly intervals between June andSeptember in 1956 and May and October in 1957. To minimize theeffects of ontogenetic drift pots containing plants of a similarmorphological status were selected from batches sown every fewdays. At the beginning and end of each experiment replicatedand paired pots were harvested and the dry weights of the leaves,stems, and roots together with the leaf areas determined. Fromthese data weekly values for net assimilation rate, leaf-arearatio (ratio of leaf area to plant weight), and relative growthrate were calculated. Simultaneously, records were kept of the diurnal changes inair temperature and of light energy by means of an integratingphotometer. Multiple regressions linking light and temperature with netassimilation rate, leaf-area ratio, and relative growth ratewere calculated separately for each year. A significant ‘time-of-season’trend was largely eliminated by including an additional variable,the initial leaf-area ratio. In the individual regressions thevariance accounted for was very high, ranging from 75 to 97per cent. The results demonstrated that for both species the net assimilationrate and relative growth rate were positively dependent on lightand temperature. The leaf-area ratio of both species was negativelyaffected by light, but only for V. faba was there a positiverelationship between the leaf-area ratio and temperature. H.annuus grew faster than V. faba during the major part of theseason, largely because of its higher leaf-area ratio. The results are compared with prior investigations in Englandand elsewhere.     

Effects of Carbon Dioxide Concentration on the Growth of Callistephus chinensis cultivar Johannistag

Carbon dioxide enrichment to 600 ppm increased the amount ofdry matter produced by Callistephus chinensis plants in growthcabinets with negligible mutual shading over a period of 18weeks. Further enrichment to 900 ppm showed smaller and morevariable increases. These effects were the result of a higherunit leaf rate of the treated plants. The direct effect on unitleaf rate was partly offset by a reduction in leaf-area ratio,and this was due almost entirely to the effect on specific leafarea with hardly any effect on leaf-weight ratio. Carbon dioxideaccelerated flower development by about a week at 600 ppm andsomewhat less at 900 ppm. The proportion of the total plantweight in the form of flowers showed a similar trend with timein all treatments and the relationship between flower-weightratio and dry-matter content of flowers was likewise similarfor all treatments, with the highest dry-matter contents ofabout 19 per cent associated with the highest flower-weightratios of about 0.44 for mature flowers. Carbon dioxide enrichmentsignificantly increased the dry-matter content of leaves. Theefficiency of energy conversion based on incident light anda twenty-four-hour cycle of 8 h light and 16 h dark for smallplants of 140–300 mg total dry weight (leaf areas of 50–120cm²) was about 4.7 per cent for the 325 ppm treatment, 6.3 percent for 600 ppm, and 5.5 per cent for 900 ppm. By referenceto some further experiments on the growth of C. chinensis cultivarJohannistag in glasshouse conditions, considerable adaptiveresponse to high and low light intensity was also demonstrated.     

The Variation in Response to Light Intensity and Carbon Dioxide Concentration Shown by Two Cultivars of Chrysanthemum morifolium Grown in Controlled Environments at Two Times of Year

The growth of the cultivar Golden Princess Anne (G.P.A.) wasstudied in controlled announcement cabinets in a range of lightconditions (125–375 J cm^–2 8-h day^–1) andcarbon dioxide concentrations (325–1500 ppm) in all combinationsPlants obtained in January and grown from January to April showedgreater final total dry weight and flower dry weight at bothhigher light intensity and higher carbon dioxide concentrationwith a strong positive interaction between them, whereas plantsobtained in September and grown from September to December didnot respond much to increased carbon dioxide concentration andthere was only a small positive interaction with light intensity.The plants grown from January to April had larger final leafareas, larger mean leaf-area ratios due mainly to larger specificleaf areas, and higher mean specific leaf-water contents comparedwith September–December plants. Despite the differencein specific leaf-water content, leaf area was almost the samelinear function of absolute leaf-water content at both timesof year. The other vegetative parts also had higher specificwater contents throughout the January–April experimentand the lateral branches were longer when compared with thecorresponding values for September–December Flower developmentwas slightly faster in September–December and the plantsbore on average one flowering branch less compared with January–Aprilplants. Plants in the lower light and carbon dioxide conditions hadlower unit leaf rates, but for plants of similar total dry weightthe effects of this on dry-matter increment were partially offsetby larger leaf areas at both times of year. The January–Aprilplants had greater leaf areas than September–Decemberplants of similar unit leaf rate and total dry weight. The cultivar Bright Golden Anne (B.G.A ) showed effects whichwere in the same direction but smaller in magnitude, tendingto diminish the differences between the times of year For example,the positive interaction in total plant dry weight was smallerin January–April compared with G P A , but larger in September–December.Leaf area, leaf-area ratio, specific leaf area, specific watercontent of leaf, stem, and root, and lateral branch length,were all larger for B G A in corresponding treatment-combinationsin two January–April experiments than in a September–Decemberone, although the difference between the times of year was smallerthan for G.P.A except for leaf area which was relatively butnot absolutely smaller Dry-matter increment and leaf area showedan inverse relationship for plants of the same total dry weight,as in G P A. In January–April B G.A plants of similarunit leaf rate and total dry weight also had greater leaf areasthan in September–December but the differences were notso large as for G.P.A Total dry-matter production was slightlygreater for B.G.A. in January–April and considerably greaterin September–December compared with G P A , and at bothtimes of year B.G.A. was more leafy, with higher specific watercontents for the vegetative parts. It was not possible to determine the cause of the differencesin growth obtained at the two times of year. It could have arisenbefore the cuttings were removed from the stock plants, duringpropagation, or during the course of the experiments in thegrowth cabinets.     

The Effects of Light Intensity and Carbon Dioxide Concentration on the Growth of Chrysanthemum morifolium cv. Bright Golden Anne

Rooted cuttings were grown in controlled-environment cabinetsat daily visible light totals of 31, 63, 125, and 250 J cm^–28-h day^–1 and carbon dioxide concentrations of 325 and600 ppm. The experiment was repeated on another occasion withthe inclusion of a further carbon dioxide level of 900 ppm.A 5-h tungsten night break was used in the first week to delayflower initiation The plants in the various treatment combinationswere sampled by frequent small harvests for leaf area and freshand dry weights of leaf, stem, root, and flower, and also forvarious morphological features. Other growth measures were obtainedby manipulation of the primary data, including the fitting ofprogress curves. Plants were respaced at intervals to minimizemutual shading. There was an increase in total dry-matter production with increasinglight and carbon dioxide, with a small positive interactionbetween them. Plants in one experiment had a somewhat higherunit leaf rate and a lower leaf-area ratio, the latter beingdue to a slightly smaller leaf-weight ratio. The effects ofadditional carbon dioxide were largely accounted for by increasedphotosynthesis. Although there were substantial differencesin specific leaf area between treatment combinations withineach experiment, the leaf-weight ratio was little altered inthe period of vegetative growth. The inverse relationship betweenspecific leaf area and unit leaf rate showed a very similartrend for all combinations of light and carbon dioxide concentration.Leaf area was a linear function of absolute leaf water contentfor all treatment combinations within an experiment, but therewas a small significant difference between occasions. Flower development was extremely delayed in the lowest lightlevel and substantially delayed at the next higher level. Thenumber of leaves below the flower decreased with increasinglight level Flower weight increased with increasing light above63 J cm^–2 8-h day^–1 and with increasing carbon dioxideconcentration, there being a positive interaction between them The initial weight and leaf area of cuttings differed for thetwo experiments, and although the results on the two occasionswere in the same direction, their magnitudes were different.Some of the discrepancy was eliminated by expressing the variousgrowth measures as functions of plant dry weight, but therewas evidently a difference in the potential for growth of thetwo batches of cuttings. The plants which were initially smallerhad a higher average unit leaf rate which, due to a higher leafwater content, was not offset by a lower leaf area ratio.     

Growth and Dry-weight Distribution in Callistephus chinensis as Influenced by Lighting Treatment

Plants of two cultivars of Callistephus chinensis (Queen ofthe Market and Johannistag) were grown in 8 h of daylight perday with one of the following treatments given during the 16h dark period: (a) darkness—‘uninterrupted night’,(b) I h of light in the middle of the dark period—a ‘nightbreak’, (c) I min of light in every hour of the dark period—‘cycliclighting’, (d) light throughout—‘continuouslight’. The plants receiving uninterrupted dark periods remained compactand rosetted in habit with small leaves, while leaf expansion,stem extension, and flower initiation were promoted in all threeillumination treatments (b, c, d). Although these three treatmentsproduced similar increases in leaf area, continuous light wasthe most effective for the promotion of both stem growth andflower initiation while cyclic lighting was generally more effectivethan a I-h night break. Continuous light also caused more dry matter to be divertedto stems at any given vegetative dry weight and it was shownthat the stem weight ratio of both varieties was correlatedwith stem length.     

The effect of light intensity on stomatal frequency in leaves ofIris hollandica hort., var. wedgwood

Stomatal frequency in leaves of plants of Iris hollandica, cultivated under four light intensities, about 12%, 37%, 75% and 100% of natural daylight, was studied. The stomatal frequency decreases with lower light intensities. The gradients of stomatal frequency of successive leaves show an increase up to the third or fourth leaf with a subsequent decrease of values up to the highest leaf. The total number of stomata per leaf is not constant; differences exist in successive leaves of a shoot as well as among the leaves of plants growing under various light intensities. The character of gradients was changed at lower light intensities. The largest reaction to the light intensity appeared in leaves with higher stomatal frequency. The gradients in individual leaves depend on the insertion of the leaf. Both the shape and the slope of these gradients are influenced by the light intensity.     

Assimilation Rate and Growth of Lolium Populations in the Glasshouse in Contrasting Light Intensities

Using growth-analysis techniques, the variation in relativegrowth-rate (RGR) and its components, net assimilation rate(NAR), and leaf-area ratio (LAR), was examined in 18 populationsof L. perenne, six of L. multiflorum, and two hybrid cultivarsfrom contrasting climatic and agronomic origins, grown at lowand high light intensities in the glasshouse. Significant differences between populations were found for RGR,NAR, and LAR at both light intensities. At both intensitiesthe annual or biennial multiflorum group had a greater LAR anda lower specific leaf weight and chlorophyll content than theperennial perenne group. At the low intensity this was compensatedby a greater NAR in the perenne group, with no resultant differencein RGR. At the high intensity there was no difference betweenthe groups in NAR, and hence a greater RGR in the multiflorumgroup. Within the perenne and multiflorum groups, at both light intensities,the variation between populations in RGR was based on differencesin NAR rather than in LAR. There was no regular correlationof NAR with either specific leaf weight, or chlorophyll contentat either light intensity, though at low light intensity itwas significantly correlated with shoot-root ratio.     

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.     

Differences in the Interacting Effects of Light and Temperature on Growth of Four Species in the Vegetative Phase

A comparative study, employing the concepts of growth analysis,has been made of the varying responses in the early vegetativephase of Gossypium hirsutum, Helianthus annuus, Phaseolus vulgaris,and Zea mays to combinations of light intensity (1.08, 2.16,3.24, 4.32, and 5.4 x 10⁴ lx—photoperiod 14 h) and constantdiurnal air temperatures (10, 15, 20, 25, 30, and 35 °C).Depending on the combination of treatments, the temperatureof the internal tissues departed from air temperature by 6.9to 1.4 °C: so only the internal temperatures are cited here. For each species there are complex interactions between theeffects of light and temperature on the net assimilation rate,the leaf-area ratio, and the relative growth-rates of plantweight and leaf area. The magnitude of the changes induced bythe two factors vary both with the growth component and thespecies. The temperature responses are maximal up to 20–5°C while at the highest temperatures they may be negative.The temperature coefficients for leaf-area ratio are consistentlyless than those of the other three components: here betweenspecies the coefficients over 10–20 °C vary by a factorof 9.6, 5.4, and 5.1 for the rates of gain in plant weight andleaf area and the net assimilation rate, while the orderingwithin each growth component is species dependent. Under conditions of optimal temperature the relative growth-rateand net assimilation rate progressively increase, accordingto the species, up to either 4.32 or 5.4x 10⁴ lx. The leaf-arearatio is always largest at the lowest intensity. The level oflight at which the rate of gain in leaf area reaches a maximumranges from 2.16x 10⁴ lx for Phaseolus to between 4.32 and 5.40x10⁴ lx for Gossypium. The highest relative growth-rate and net assimilation rate ofHelianthus exceed those of Zea substantially. Indeed the maximalassimilation rate for Helianthus of 2.10 g dm^–2 week^–1is the highest ever recorded under field or controlled conditions.Possible reasons for this reversal of the photosynthetic potentialsof the two species observed by previous workers are discussed.     

The Relationship between Sugar and Amino Acid Export to the Phloem in Young Wheat Plants

The relationship between amino acid and sugar export to thephloem was studied in young wheat plants (Triticum aestivumL. ‘Pro-INTA, Isla Verde’) using the EDTA-phloemcollection technique. Plants grown with a 16 h photoperiod showeda rapid decrease in the concentration of sugars and amino acidsin the phloem exudate from the beginning of the dark period.When plants grown with a 16 h photoperiod were kept in the darkfor longer than 8 h the free amino acid content in leaves andexudate (on a dry weight basis) increased continually throughoutthe 72 h of darkness. During the first 24 h of darkness thesugars in the phloem exudate decreased to 30% of the initialvalue, and returned to the control level when plants were returnedto light. When plants grown under low light intensity for 10d were transferred to high light intensity, they showed an increasein leaf sugar content (dry weight basis) after 3 d but therewere no differences in leaf free amino acid content (dry weightbasis) compared to low-light plants. The sugar concentrationin the phloem exudate was increased by higher light intensities,but there was no difference in the amino acid concentrationof the phloem exudate, and thus the amino acid:sugar ratio inthe phloem decreased in the high-light plants. The present resultssuggest that amino acids can be exported to the phloem independentlyof the export of sugars. Copyright 1999 Annals of Botany Company Sugar exudation, amino acid transport, nitrogen, phloem, transport, wheat, Triticum aestivum L.     

Diurnal courses and factorial dependencies of leaf conductance and transpiration of differently potassium fertilized and watered field grown barley plants

During the grain filling period we followed diurnal courses in leaf water potential (ψ₁), leaf osmotic potential (ψ_π), transpiration (E), leaf conductance to water vapour transfer (g) and microclimatic parameters in field-grown spring barley (Hordeum distichum L. cv. Gunnar). The barley crop was grown on a coarse textured sandy soil at low (50 kg ha⁻¹) or high (200 kg ha⁻¹) levels of potassium applied as KCl. The investigation was undertaken at full irrigation or under drought. Drought was imposed at the beginning of the grain filling period. Leaf conductance and rate of transpiration were higher in the flag leaf than in the leaves of lower insertion. The rate of transpiration of the awns on a dry weight basis was of similar magnitude to that of the flag leaves. On clear days the rate of transpiration of fully watered barley plants was at a high level during most part of the day. The transpiration only decreased at low light intensities. The rate of transpiration was high despite leaf water potentials falling to rather low values due to high evaporative demands. In water stressed plants transpiration decreased and midday depression of transpiration occurred. Normally, daily accumulated transpirational water loss was lower in high K leaves than in low K leaves and generally the bulk water relations of the leaves were more favourable in high K plants than in low K plants. The factorial dependency of the flag leaf conductances on leaf water potential, light intensity, leaf temperature, and leaf-to-air water vapour concentration difference (ΔW) was analysed from a set of field data. From these data, similar sets of microclimatic conditions were classified, and dependencies of leaf conductance on the various environmental parameters were ascertained. The resulting mathematical functions were combined in an empirical simulation model. The results of the model were tested against other sets of measured data. Deviations between measured and predicted leaf conductance occurred at low light intensities. In the flag leaf, water potentials below-1.6 MPa reduced the stomatal apertures and determined the upper limit of leaf conductance. In leaves of lower insertion level conductances were reduced already at higher leaf water potentials. Leaf conductance was increased hyperbolically as photosynthetic active radiation (PAR) increased from darkness to full light. Leaf conductance as a function of leaf temperature followed an optimum curve which in the model was replaced by two linear regression lines intersecting at the optimum temperature of 23.4°C. Increasing leaf-to-air water vapour concentration difference caused a linear decrease in leaf conductance. Leaf conductances became slightly more reduced by lowered water potentials in the low K plants. Stomatal closure in response to a temperature change away from the optimum was more sensitive in high K plants, and also the decrease in leaf conductance under the influence of lowered ambient humidity proceeded with a higher sensitivity in high K plants. Thus, under conditions which favoured high conductances increase of evaporative demand caused an about 10% larger decrease in leaf conductance in the high K plants than in the low K plants. Stomatal sizes and density in the flag leaves differed between low and high K plants. In plants with partially open stomata, leaf conductance, calculated from stomatal pore dimensions, was up to 10% lower in the high K plants than in the low K plants. A similar reduction in leaf conductance in high K plants was measured porometrically. It was concluded that the beneficial effect of K supply on water use efficiency reported in former studies primarily resulted from altered stomatal sizes and densities.     

Physiological and Ecological Studies in the Analysis of Plant Environment: XII. The Role of the Light Factor in Limiting Growth

Previous investigations in southern England on twenty-two herbaceousspecies have demonstrated that for widely spaced plants thediurnal solar radiation limits the net assimilation rate ofall species and restricts the relative growth rate of many.In examining how far these limitations apply to other environmentsit is now shown that in the subtropics and tropics the levelsof net assimilation rate and relative growth rate can greatlyexceed those so far recorded for cool temperate regions, andthese differences are attributed to the higher insolation andtemperatures. From a variety of evidence it is concluded that as the distancebetween plants is reduced 8O the net assimilation rate is progressivelydiminished even in regions of high insolation through the enhancedmutual shading. In consequence levels of light which may besupra-optimal for relatively isolated individuals may yet limitthe dry-matter production of a dense population. There is anoptimal ratio of leaf area to ground surface (leaf-area index)for the maximal exploitation of the incoming radiation in carbonfixation by the population and this optimum will vary with thespecies and the light intensity. Where other environmental factorsare favourable, light may limit dry-matter production everywhere. On an annual basis dry-matter production will be dependent ontwo components—the length of the ‘growing season’and the period over which the leaf-area index remains optimal.In the tropics the highest annual rate of production so farrecorded is 7⁸ tonnes/hect. produced by Saccharum officinarumandin north-east Europe 23.5 tonnes by Fagus sylvatica. Over shortperiods the rate of dry-matter production can attain 3⁸g./m.²/dayand the utilization of solar energy can be as high as 4.2 percent., or 9.5 per cent, for the range 4, 000–7, 000 A. Although information on the productivity of natural communitiesis still ex-ceedingly scanty, an attempt has been made to interpretthe general pattern in terms of the length of the growing season,the level of solar radiation, the magni-tude of the leaf-areaindex of the whole community, and the period over which theleaf canopy remains green. It is postulated that in any regionthe vegetation reaches a dynamic equilibrium when there is themaximum exploitation of the incoming radiation to produce thegreatest production of dry matter.     

Photochemical and photophosphorylation activities of chloroplasts and leaf mesostructure in Chinese cabbage plants grown under illumination with light-emitting diodes

Leaf mesostructure, photochemical activity, and chloroplast photophosphorylation (PP) in the fourth true leaf of 28-day-old Chinese cabbage (Brassica chinensis L.) plants were investigated. Plants were grown under a light source based on red (650 nm) and blue (470 nm) light-emitting diodes (LED) with red/blue photon flux ratio of 7: 1 and under illumination with high-pressure sodium lamp (HPSL) at photon flux densities of 391 ± 24 μmol/(m² s) (“normal irradiance”) and 107 ± 9 μmol/(m² s) (“low irradiance”) in photosynthetically active range. At normal irradiance, the leaf area in plants grown under HPSL was twofold higher than in LED-illuminated plants; other parameters of leaf mesostructure were little affected by spectral quality of incident light. The lowering of growth irradiance reduced the majority of leaf mesostructure parameters in plants grown under illumination with HPSL, whereas in LED-illuminated plants the lowered irradiance reduced only specific leaf weight but increased the leaf thickness and dimensions of mesophyll cells and chloroplasts. The photochemical activity of isolated chloroplasts was almost independent of growth irradiance and light spectral quality. Light quality and intensity used for plant growing had a considerable impact on PP in chloroplasts. At normal light intensity, the highest activity of noncyclic PP in chloroplasts was observed for plants grown under HPSL; at low light intensity the highest rates of PP were noted for plants grown under LED. The P/2e⁻ ratio, which characterizes the degree of PP coupling to electron transport in the chloroplast electron transport chain, showed a similar pattern. Thus, the narrow-band spectrum of the light source had little influence on leaf mesostructure and electron transport rates. However, this spectrum significantly affected the chloroplast PP activity. The PP patterns at low and normal light intensities were opposite for plants grown under LED and HPSL light sources. We suppose that growing plants under LED array at normal light intensity disturbed the chloroplast coupling system, thus preventing the effective use of light energy for ATP synthesis. At low light intensity, chloroplast PP activity was significantly higher under LED illumination, but plant growth was suppressed because of impaired adaptation to low light intensity.     

A Comparison of the Effects of Light Intensity and Duration on Chrysanthemum morifolium cv. Bright Golden Anne in Controlled Environments: I. Growth Analysis

Rooted cuttings were grown at a constant temperature of 15.6°C in two light intensities (22 and 33 J m–² sec–¹)in either 8- or 12-h days. The combination of the lower lightand the longer day gave the same daily light total as the reversecombination. Although departures from reciprocity were verysmall, total dry weight, specific leaf area, leaf area ratio,leaf weight ratio, and specific water content of stems weresignificantly greater for the lower light-longer day treatment.Only for specific water content of roots was the value for thelower light-shorter day treatment significantly smaller. Unitleaf rate was a linear function of light total, the two treatmentswith the same daily total being almost identical. In comparisonwith previous experiments, the plants had smaller leaf areasrelative to their rate of dry-matter increment. This is attributedto the lower day temperature. Flowering was earliest in thehigher light-shorter day combination, but surprisingly in thelonger day the higher intensity did not result in earlier floweringthan the lower intensity. The relationship between flower weightratio and flower development was the same as for other experimentsat different temperatures.     

Effect of temporary changes in light intensity on carbon transport, partitioning and respiratory loss in young tomato seedlings raised under different light intensities

Tomato plants were grown under light intensities of 36 or 90 W m⁻² [photosynthetically active radiation (PAR)], and then the light intensity was changed to 36, 90 or 180 W m⁻² for 8 h to investigate the effect of temporary changes in light intensity on the carbon budget of photoassimilates from the third leaf using a ¹⁴CO₂ steady-state feeding method. In the plants that were raised under 90 W m⁻², the photosynthetic rate increased when the light intensity was increased to 180 W m⁻², whereas no increase occurred in the plants that were raised under 36 W m⁻². Although the total amount of carbon fixed during the 8-h light period showed a large difference between plants grown at the two initial light intensities, the proportion of carbon exported during the light period did not differ apparently, irrespective of the change in light intensity. However, the amount of carbon exported during the time course was higher in plants that were raised under 90 W m⁻² than those raised under 36 W m⁻², irrespective of the change in light intensity. The partitioning pattern of ¹⁴C-photoassimilates was not changed by the change in light intensity, irrespective of whether the light intensity was increased or not. However, the amount of ¹⁴C-photoassimilates accumulated in each part differed according to the two initial light intensities. The carbon transport from a source leaf was also investigated through a quantitative analysis of carbon balance.     

Effects of Natural and Artificial Light in Arctic Latitudes on Long- and Short-day Plants as Revealed by Growth Analysis

The effects on growth and flowering of two short-day and twolong-day plants when grown under different conditions of illuminationare described. The plants fully investigated were Kalanchoeblossfeldiana and Xanthium pennsylvanicum and the annual varietiesof Hyoscyamus niger and Beta vulgaris. Wintex barley, Iberisumbellata, and tomato were also grown in some selected treatments.The conditions investigated comprised continuous full daylight(24 hours), full daylight for the whole of the daily photoperiodand full daylight for half the photoperiods, the other halfconsisting of either daylight reduced by shading or light fromincandescent lamps or fluorescent tubes (daylight-matching type),all of the same low intensity. Two lengths of photoperiod wereused for each species, one nearly optimal for flowering, theother closer to the critical day-length; and the order of thelow and high light treatments was varied. These factors werecombined factorially. Data were collected (or derived) for the following characteristics,though not always for all the species grown: dry weights, leafareas, heights, water contents, epidermal cell sizes, net assimilationrates, times to flowering, leaf-number increments until flowering,numbers of inflorescences, stomatal apertures, and leaf postures. Among other effects, the data revealed that in all four speciesinvestigated the adverse effects on over-all growth to be expectedfrom reduction of the daily photoperiod or of the intensityof illumination are in fact minimized. This compensation waseffected mainly by large increases in leaf areas, even thoughin all cases half the daily photoperiod consisted of full daylight.There are indications that increased efficiencies (net assimilationrates) may also have been involved. The leaf-area increasesappear to have been due to increased cell size rather than cellnumber and a close positive correlation with water content wasfound. The most striking among the effects on flowering was the failureof sugar-beet to bolt when half of its photoperiod (totals of20 and 14 hours) consisted of light from fluorescent lamps.The flowering of barley and Hyoscyamus was also delayed considerablyunder these conditions. The deficiency of red in the spectrumof the fluorescent light is believed to have been the cause.By contrast, the flowering of Iberis, a crucifer, was not affected.