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.     

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: XI. A Further Assessment of the Influence of Shading on the Growth of Different Species in the Vegetative Phase

In a further analysis of the effects of varying light intensityon growth and development in the vegetative phase the reactionsof thirteen herbaceous species have been recorded. In some experimentsthe degree of shading has been extended to 0.055 daylight, alevel near or below the compensation point. For Lathyrus maritimus, Trifolium pratense, and Vicia faba,the net assimilation rate is directly related to the logarithmof the light intensity, but for Helianthus annuus, T. repent,T. hybridum, Medicago sativa, Phaseolus multiflorus, and Loliummultiflorum the relationship, though curvilinear, is not logarithmic.It is concluded that for all species the assimilation rate ofunshaded plants was limited by light even though in high summerthe recorded light energy between 4,000–7,000 A averaged1,900–2,200 foot-candles. For all these species between daylight and 0.12 daylight theleaf-area ratio rises as the intensity decreases and in generalthe trend is logarithmic. When the degree of shading is increasedto 0.055 daylight the logarithmic relationship still holds forL. maritimus and V. faba though this level is below the compensationpoint. For other species, such as P. multiflorus andH. annuusthe trend may be reversed below 0.12 daylight and the ratiothen falls. When the light intensity is reduced from daylight to 0.5 daylight,then for the species already cited and for Lolium perenne, Phleumpratcnse, and Festuca pratensis the relative growth-rate isinvariably depressed. At 0.055 daylight the relative growth-ratenever exceeded 1 per cent. per day. For L. perennet, P. pratense,and Dactylis glomerata the reactions to shading of ‘hay’and ‘grazing’ strains were different. The ecological and physiological implications of these findingsare discussed.     

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.     

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.     

Gas Flow Measurement in Fermentations: An Inexpensive System with Analogue Output for Recording

During the fermentation of North Sea Gas to single cell protein in a gas-recycle system (Spivey 1973) there was a need to measure continuously low gas flow rates and to obtain a recorder signal proportional to the instantaneous flow rate. Commercial equipment, such as differential pressure transmitters linked to a pneumatic recorder, was expensive and not readily modified for the low flow rates of laboratory fermentations.     

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.     

Physiological Studies in Plant Nutrition: XVI. The Mineral Nutrition of Bracken

Part I of this paper describes culture methods for the productionof large numbers of bracken sporelings of uniform size. Observationson the growth of the bracken plant and results of experimentson the mineral nutrition of the prothallial and sporophyte generationsare presented. The presentation of further growth data and afull analysis of the results is deferred to the second partof this paper.     

An Analysis of Growth of Oil Palm Seedlings in Full Daylight and in Shade

Growth of seedling oil palms (Elaeis guineensis), in full daylightand under three levels of shade, was studied using growth analysistechniques. In full day-light, net assimilation rates (E_A) betweeno.15 and 0.32 g./dm.²/week were obtained associated with lowleaf-area ratios (F) giving relative growth-rates (R_W) rangingfromI I.8 to 3.2 per cent, per day. There were no indicationsof seasonal differences within the small range of values found. The plants take about 90 days to adapt to shade conditions becausethe mean plastochron is 24 days, and shading effects are beststudied on plants grown since germination under the shade treatments.Very different pictures of response to shade were obtained usingplants grown initially in full light followed by 90 days' shadebefore sampling compared with plants grown under shade sincegermination. In the latter, except at the lowest light levelused, I I.I per cent, of full day-light, there was very littleeffect of light on E_A or R_w, although the F values decreasedas light increased. Extrapolation of the F values to the extinctionpoint gave values similar to those obtained in another experimenton the effect of a number of shade levels on F. The physiological and ecological implications of these findings,particularly the low growth-rates and shade tolerance, are discussed.     

An Ecological Risk Assessment of Phenol in the Aquatic Environment

A probabilistic ecological risk assessment of phenol was undertaken to determine the risks posed to biota as a result of phenol release to the Canadian environment. A three-tiered approach was used to estimate risks, with progressively more realistic assumptions being applied at each tier. In Canada, the major sources of phenol are municipal wastewater treatment plants, pulp, paper and wood products mills, steel and metal products facilities and refineries. Thus, the highest exposures will occur in receiving waters near these point sources, primarily due to the short half-life of phenol in the aquatic environment. Sensitive aquatic organisms include salmonids (e.g., rainbow trout Oncorhynchus mykiss) and amphibians (e.g., leopard frog Rana pipiens). The results of the risk assessment indicate that species are exposed to elevated levels of phenol near point sources, but these levels represent only a minor risk to aquatic biota.     

An Ecological Risk Assessment of Ammonia in the Aquatic Environment

Ammonia is released in the environment by many industries and other human activities. The major quantifiable sources of ammonia released to aquatic ecosystems across Canada are municipal wastewater treatment plants, at an estimated total quantity of 62,000 tonnes per year. Given the sources of ammonia releases in the environment and the properties of the substance, terrestrial plants and aquatic organisms are potential risk targets. A tiered assessment approach has been used to determine the ecological risk in the aquatic environment from ammonia released in municipal wastewater effluents. The results obtained for two case studies with the probabilistic risk analysis used in the highest tier support the conclusion that the conditions encountered in these two locations can lead to ammonia concentrations capable of producing an adverse ecological impact.     

Studies of the Surfaces of Desert Plant Seeds: II. Ecological Adaptations of the Seeds of Blepharis persica

A study was made of the role of the hydrated mucilage whicharises from the integumentary hairs of Blepharis seeds and inhibitsgermination where there is an excess of water, and of the anatomicaland physiological basis for this phenomenon. It was found thatthis inhibition was not due to bacterial respiration. The appearanceof the dry mucilage in the unimbibed seeds is described fromscanning electron micrographs. The basic helical framework ofthe secondary wall thickenings and the very unusual second-orderframework of trabeculae in the integumentary hair cells aredescribed.     

Kinetic Measurements of Small Ethylene Changes in an Open System Designed for Plant Physiological Studies

Based upon the method we developed to measure ethylene in very low concentrations (as low as 0.01 ppb in the ambient atmosphere) an experimental chamber was constructed and integrated in the measuring system for plant physiological studies. Parameters influencing the accuracy of the technique are evaluated. The pressure in the plant chamber increased 0.05 atm at a flow rate of 10 1/h during ethylene trapping. At this flow rate the ethylene production per seedling is independent of the number of seedlings used. The ethylene measured per seedling is directly proportional to the length of trapping time. Under standard conditions of chamber configuration and volume very small changes in ethylene content can be detected accurately in a very short time range. Our experimental arrangement allows kinetic studies of ethylene evolution by biological objects in a qualitative and quantitative manner. All methods used heretofore are more complicated and less accurate compared to the measuring system presented here. Its versatility is demonstrated for both in vitro and in vivo studies of ethylene production by bean seedlings. The application fields of the apparatus are discussed.     

Studies in the Respiratory and Carbohydrate Metabolism of Plant Tissues3: VIII. AN INHIBITION OF RESPIRATION IN PEAS INDUCED BY 'OXYGEN POISONING'

A pressure of oxygen of 5 atmospheres has been shown to inhibitmarkedly the carbon-dioxide output of green shelled peas. Accompanyingthis decrease, an accumulation of pyruvic and ‘citric’acid has been noted, whilst a depletion in the concentrationof -ketoglutaric acid and, in some experiments, of malic acidoccurred. The results have been interpreted as indicating thathigh pressures of oxygen interfere with the metabolism of citricacid causing a ‘jamming’ of the tricarboxylic acidcycle; thus, under the conditions prevailing during the oxygentreatment, the tricarboxylic acid cycle is considered to bea major respiratory pathway in peas.     

Current-Voltage Curves for Plant Membrane Studies: A Critical Analysis of the Method

The evolution of the steady-state current-voltage (I/V) methodis followed to its present sophistication of bipolar staircase.When applied to Chara plasmalemma the resultant I/V characteristicsvary enormously depending on the physiological state of themembrane (which is, in turn, dictated by the outside conditions).This variety of response makes formulation of ‘correct’,that is artefact-free, I/V technique quite a challenge. Thetime-dependence of the clamp current in each electrophysiologicalstate is different and this must be taken into considerationwhen steady-state I/V is investigated. Clamping of the plasmalemmaalone is also necessary, as under some conditions the conductanceof the two membranes becomes similar and the effects of thetonoplast can no longer be neglected when both membranes areclamped in series. The data obtained by point clamping and spaceclamping are compared. The use of bipolar staircase introducesmore subtle artifacts: the I/V profiles can be severely misinterpreteddue to transient conductance changes arising from the excitationor the outward and inward rectifier currents. The excitationcan be blocked by temporary exposure to lanthanum ion, but theconcentration must be carefully chosen. It is necessary to optimizethe staircase parameters to allow the rectifier currents toreturn to resting level before the next staircase pulse. Finally,brief comparison of the Chara I/Vprofile to that of other cellsis included and the relevance to patch clamp studies discussed. Key words: Current-voltage characteristics, Chara, space clamp, lanthanum blockade     

An Ecological Risk Assessment of Nonylphenol and Its Ethoxylates in the Aquatic Environment

Nonylphenol ethoxylates (NPEs) are a group of surfactants that are widely used for industrial, commercial, institutional and household purposes in Canada. Ethoxylation of nonylphenol (NP) occurs upon reaction with ethylene oxide, producing NPEs, although NP is also used in the production of the antioxidant tris(nonylphenol)phosphite. NP and NPEs are not produced naturally, and the primary route of environmental exposure to NP and NPEs is via textile mill, pulp and paper mill and municipal wastewater treatment plant effluents. NPEs occur as complex mixtures and are described by the average ethoxylate chain length, which ranges from 1 to 100. The environmental fate of NPEs is strongly dependent on the effluent and, the degree and type of treatment to which the effluent is subjected. An ecological risk assessment was performed to determine if exposure to NP and NPEs results in effects on the Canadian environment, based on current use patterns. The Canadian ecological risk assessment found that adverse effects on aquatic organisms are likely, although assumptions were made with respect to appropriate dilution factors.     

Studies on the Growth in Culture of Plant Cells: VIII. THE PRODUCTION OF ETHYLENE BY SUSPENSION CULTURES OF ACER PSEUDOPLATANUS, L.

Sycamore cell suspension cultures in a synthetic medium releaseethylene; during a 24-day incubation period a single culture(initial volume 70 ml) produces c. 4 µ moles. There isa very sharp peak of ethylene production between day 10 andday 14 of culture; at the peak of production c. 2 nmoles ethyleneare released per million cells in 24 h. Evidence is presentedthat 2,4-D enhances ethylene production independently of itseffects on culture growth. Under the standard conditions of culture (250-ml Erlenmeyerflasks closed with aluminium foil and containing 70 ml cellsuspension) the concentration of ethylene in the gas phase ofthe cultures rises above 10 ppm. No evidence was obtained thatthis ethylene is inhibitory to culture growth or that a criticallevel of ethylene is necessary to initiate cell division incultures at a critically low cell density. The low rate of ethylene release by stationary phase culturesis temporarily enhanced by the addition of various solutes andfurther depressed by dilution with water.