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.     

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.     

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.     

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: 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.     

The Importance of Light Intensity for Pollen Tube Growth and Embryo Survival in Wheat x Maize Crosses

The success of Triticum aestivumxZea mays crosses, used to producewheat doubled haploids, is influenced by light intensity. Toexamine the basis for this response, pollen tube growth, embryosurvival and indicators of photosynthetic rate were measuredin two wheat cultivars (‘Karamu’ and ‘Kotuku’)crossed with maize at two irradiance levels (250 or 750 µmolm^-2s^-1, PAR). Pollen tube growth was significantly affectedby light intensity in ‘Karamu’ plants but not in‘Kotuku’ plants, despite both cultivars being pollinatedby the same maize source. The percentage of pollen tubes reachingthe cavity between the ovarian wall and integuments, or in themicropyle of ‘Karamu’ plants at high light intensity(65%) was nearly three-times greater than that at low lightintensity (22%). Thus, either low light intensity can affectthe maternal wheat plant in a way that inhibits pollen tubegrowth and/or high light intensity may promote pollen tube growthin ‘Karamu’ plants. Significant differences in ratesof electron transport in plants grown at the two light intensitiesindicated that the rate of photosynthesis may also have an effecton pollen tube growth. These results have importance for improvingthe efficiency of wheat x maize crosses and other wide cerealcrosses. Copyright 2001 Annals of Botany Company Intergeneric hybridization, light intensity, pollen tube growth, embryo survival, Triticum aestivum, wheat,Zea mays , maize     

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.     

Influence of Flooding on Net CO2Assimilation, Growth and Stem Anatomy of Annona Species

A series of experiments was conducted to assess net CO₂assimilationand growth responses to waterlogging of grafted and seedlingtrees in the genus Annona. Seedlings of A. glabra, A. muricataandA. squamosa L., and scions of ‘Gefner’ atemoya(A. squamosaxA. cherimola Mill.), ‘49-11’ (‘Gefner’atemoyaxA. reticulata L.), ‘4-5’ (‘Priestley’atemoyaxA. reticulata), A. reticulata grafted onto either A.glabra, A. reticulata orA. squamosa rootstocks were floodedfor up to 60 d. Soil anaerobiosis occurred on the third dayof flooding. Seedlings ofA. glabra and A. muricata, and thescions ‘49-11’, ‘Gefner’ atemoya, andA. reticulata grafted onto A. glabra rootstock were consideredflood tolerant based on their ability to survive and grow inflooded conditions. Scions of the normally flood-sensitive A.reticulata, ‘Gefner’ atemoya, and ‘49-11’tolerated root waterlogging when grafted onto the flood-tolerantspecies, A. glabra. In contrast, flooding of A. squamosa seedlingsand rootstocks, and A. reticulata rootstocks greatly reducedgrowth and net CO₂assimilation rates, and resulted in 20–80%tree mortality. Stem anatomical responses to long-term flooding(12 continuous months) were assessed in seedlings of A. glabraand A. muricata, and trees of ‘49-11’ grafted ontoA. glabra. Flooded trees developed hypertrophied stem lenticels,particularly in A. glabra, and enlarged xylem cells resultingin thicker stems with reduced xylem density. Flooding did notincrease air spaces in pre-existing xylem near the pith or inxylem tissue that was formed during flooding. Thus, flood tolerancedid not involve aerenchyma formation in the stem. Copyright1999 Annals of Botany Company Flood tolerance, net CO₂assimilation, photosynthesis, stem anatomy, shoot growth, anaerobiosis, Annonaceae.     

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.     

The Morphology of Protonema and Bud Formation in the Bryales

The development of protonema from spore in aseptic culture wasfollowed in species representing 14 genera and 7 orders of theBryales. In no case could a distinction be made between sharplydefined ‘chloronema’ and ‘caulonema’stages. In all species, with the exception of the anomalous Cinclidotusfontinaloides, the protonema developed a markedly heterotrichoushabit, with prostrate filaments of relatively unlimited growthand erect filaments of more restricted development. The twotypes of filament were usually distinct in cellular characteristicsas well as in the direction of growth. In some cultures a thirdtype of filament grew downwards into the medium. Shoot buds were found to occupy diverse positions in differentspecies, but the commonest position was near the base of a primaryfilament of the erect branch system. Shoot buds were formedas readily in artificial light of sufficient intensity as innatural daylight.     

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.     

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.     

Comparison of carbon-specific growth rates and rates of cellular increase of phytoplankton in large limnetic enclosures

Potential carbon-specific growth rates of phytoplankton wereestimated from a series of measurements of photosynthetic radio-carbonuptake over 4- and 24-h exposure periods in the light fieldsof three large limnetic enclosures (‘Lund Tubes’),each providing different limnological and trophic conditions.Photosynthetic behaviour and short-term, chlorophyll-specificcarbon-fixation rates conformed to well-established criteriabut, over 24 h, the net retention represented 23–82% ofthe carbon fixed during the daylight hours. Potential mean growthrates (k'_p, of the photo-autotrophic community were calculatedas the net exponential rates of daily carbon-accumulation relativeto derived, instantaneous estimates of the cell carbon-content.Apparent actual community growth rates (k'_D were calculatedas the sum of the exponential rates of change of each of themajor species present, corrected for probable rates of in situgrazing and sinking, and expressed relative to the fractionof total biomass for which they accounted. The correspondingvalues were only occasionally similar, k'_p generally exceedingK'_D by a factor of between 1 and 30 or 1 and 14, depending uponthe carbon:chlorophyll ratio used. The ratio, K'_p/K'_D was foundto vary inversely both to k'_D and to k_n, the net rate of changein phytoplankton biomass, suggesting that measured carbon fixationrates merely represent a capacity for cellular increase which,owing to other likely limitations upon growth, is seldom realized.Apparent rates of loss of whole cells do not account for theloss of carbon; that the ‘unaccounted’ loss rates(K'_p–K'_D varied in direct proportion to K'_p (i.e., losseswere least when chlorophyll-specific photosynthetic productivitywas itself limited) is best explained by physiological voidingof excess carbon (for instance, by respiration, photorespiration,excretion) prior to the formation of new cells.     

Diverse Responses of Maple Saplings to Forest Light Regimes

Seedlings of 11 species of forest maples (AcerL.) were grownoutdoors from budburst to senescence under three light regimes:‘gap centre under clear skies’ (approx. 20% opensky irradiance; red:far-red ratio=1.12); ‘gap centre undercloudy skies’ (1.5%, ratio=1.03); and ‘gap edge’(2.5%, ratio=0.6). Seedlings grown under the gap centre (clearsky) regime had significantly greater height growth, greaterspecific leaf mass, higher root:shoot ratio, greater investmentin roots, higher leaf nitrogen concentrations, greater chlorophylla:bratio,lower photosynthetic rates under dim light, higher maximum photosyntheticrate, higher stomatal conductance, and lower leaf internal CO₂concentrationscompared with those grown in either gap edge or gap centre (cloudy)regimes. Responses to the gap edgevs.gap centre (cloudy) treatmentsdiffer little, suggesting that shade acclimation in forest mapleseedlings is mainly a response to light intensity rather thanspectral quality. The ubiquitous and, except for leaf internalCO₂concentration, highly significant interspecific variationin traits was broad-ranging and continuous. These results suggestthat (1) the responses to light quality found in shade intolerantherbaceous and woody species growing in more open habitats maynot have a selective advantage in seedlings of shade tolerantforest trees, and (2) the adaptive plastic response to understoreyvs.gapenvironments in forest maples, which is qualitatively consistentacross species, is founded on co-ordinated, small shifts insets of functionally inter-related traits.Copyright 1998 Annalsof Botany Company Acer,forest gap heterogeneity, plasticity, specific leaf mass, photosynthesis, leaf chlorophyll, nitrogen, stomatal density, root growth, root:shoot ratio, growth form.     

Plant allometry, leaf nitrogen and phosphorus stoichiometry, and interspecific trends in annual growth rates

• Background Life forms as diverse as unicellular algae,zooplankton, vascular plants, and mammals appear to obey quarter-powerscaling rules. Among the most famous of these rules is Kleiber's(i.e. basal metabolic rates scale as the three-quarters powerof body mass), which has a botanical analogue (i.e. annual plantgrowth rates scale as the three-quarters power of total bodymass). Numerous theories have tried to explain why these rulesexist, but each has been heavily criticized either on conceptualor empirical grounds. • N,P-Stoichiometry Recent models predicting growth rateson the basis of how total cell, tissue, or organism nitrogenand phosphorus are allocated, respectively, to protein and rRNAcontents may provide the answer, particularly in light of theobservation that annual plant growth rates scale linearly withrespect to standing leaf mass and that total leaf mass scalesisometrically with respect to nitrogen but as the three-quarterspower of leaf phosphorus. For example, when these relationshipsare juxtaposed with other allometric trends, a simple N,P-stoichiometricmodel successfully predicts the relative growth rates of 131diverse C₃ and C₄ species. • Conclusions The melding of allometric and N,P-stoichiometrictheoretical insights provides a robust modelling approach thatconceptually links the subcellular ‘machinery’ ofprotein/ribosomal metabolism to observed growth rates of uni-and multicellular organisms. Because the operation of this ‘machinery’is basic to the biology of all life forms, its allometry mayprovide a mechanistic explanation for the apparent ubiquityof quarter-power scaling rules.     

Integrated Analysis of Growth and Light Interception in Winter Lettuce II. Differences between Cultivars

‘Integrated growth analysis’ (emphasizing aspectsof crop and plant structure) and ‘light conversion analysis’(stressing the efficiency of interception and photosyntheticconversion of light) have been used to investigate the wintergrowth of different cultivars of butterhead and crisphead typesof glasshouse lettuce. Measurements from ‘Ambassador’ (large-framed butterhead),‘Renate’ (medium-sized butterhead) and ‘Cristallo’(crisphead) were made, statistical progressions were fittedto the primary data and hence estimates of all the analyticalcomponents were derived. Curves for crop growth rate, like those for most other components,followed a generally similar pattern for all three cultivars.In integrated growth analysis, the biomass curve for Ambassadorlay above the curves for the other cultivars. The weight advantagewas initially 60 per cent and it persisted with only a smallreduction (to 40 per cent) until the final harvest. Relativegrowth rate varied little between cultivars because differencesobserved in leaf area ratio were complementary to those seenin net assimilation rate. In light conversion analysis, differences in light interceptingefficiency between cultivars were not statistically significant,though Ambassador attained full interception 4 days earlierthan Renate and 6 days earlier than Cristallo. Differences inlight utilizing efficiency were small and non-significant exceptduring the post-rosette stage when the value for Renate waslower than that of either Ambassador or Cristallo. Deviationsaround the fitted curves were correlated with fluctuations inthe light regime. An assessment is made of the utility and limitations of thetwo procedures. It is concluded that both approaches can assistin analyzing the rate of dry matter production in crops or plantstands. Integrated growth analysis is advantageous when theneed arises to treat individual and population-based attributessimultaneously, while light conversion analysis provides a meansof explicitly incorporating the primary environmental variableinfluencing growth. Lactuca sativa L., lettuce cultivars, growth analysis, crop growth rate, relative growth rate, light interception, light utilization     

Seasonal patterns of growth, dehydrins and water-soluble carbohydrates in genotypes of Dactylis glomerata varying in summer dormancy

• Background and Aims Summer dormancy in perennial grasseshas been studied inadequately, despite its potential to enhanceplant survival and persistence in Mediterranean areas. The aimof the present work was to characterize summer dormancy anddehydration tolerance in two cultivars of Dactylis glomerata(dormant ‘Kasbah’, non-dormant ‘Oasis’)and their hybrid using physiological indicators associated withthese traits. • Methods Dehydration tolerance was assessed in a glasshouseexperiment, while seasonal metabolic changes which produce putativeprotectants for drought, such as carbohydrates and dehydrinsthat might be associated with summer dormancy, were analysedin the field. • Key Results The genotypes differed in their ability tosurvive increasing soil water deficit: lethal soil water potential(_s) was –3·4 MPa for ‘Kasbah’ (althoughnon-dormant), –1·3 MPa for ‘Oasis’,and –1·6 MPa for their hybrid. In contrast, lethalwater content of apices was similar for all genotypes (approx.0·45 g H₂O g d. wt^–1), and hence the greater survivalof ‘Kasbah’ can be ascribed to better drought avoidancerather than dehydration tolerance. In autumn-sown plants, ‘Kasbah’had greatest dormancy, the hybrid was intermediate and ‘Oasis’had none. The more dormant the genotype, the lower the metabolicactivity during summer, and the earlier the activity declinedin spring. Decreased monosaccharide content was an early indicatorof dormancy induction. Accumulation of dehydrins did not correlatewith stress tolerance, but dehydrin content was a function ofthe water status of the tissues, irrespective of the soil moisture.A protein of approx. 55 kDa occurred in leaf bases of the mostdormant cultivar even in winter. • Conclusions Drought avoidance and summer dormancy arecorrelated but can be independently expressed. These traitsare heritable, allowing selection in breeding programmes.     

Rates of Dry-matter Production in Some Tropical Forest-tree Seedlings

An analysis of growth in full daylight of four tropical forest-treeseedlings and Helianthus was made with the view of ascertainingwhether woody plants in the tropics have a lower net assimilationrate than herbaceous plants as has been demonstrated in otherenvironments. Significant differences in net assimilation ratewere found between Helianthus and three of the woody speciestested: Chlorophora excelsa, Musanga cecropioides, and Terminaliaivorensis; the fourth woody plant, Ceiba pentandra, had an unusuallyhigh net assimilation rate which, in one experiment, did notdiffer significantly from the value found for Helianthus. The data are discussed in relation to the general problems ofcomparative growth-rates of herbaceous and woody plants andof factors contributing to differences in forest productivitybetween climates.     

The Effect of Light Intensity and Relative Humidity on Growth Rate and Root Respiration of Plantago lanceolata and Zea mays

Plants of Plantago lanceolata L. and Zea mays L., cv. ‘Campo’were grown at two levels of light intensity. Especially in theroots, the rate of dry matter accumulation decreased at lowlight intensity. The carbohydrate content of both roots andshoots of P. lanceolata was not affected by light intensity.The relative contribution of SHAM¹-sensitive respiration, thealternative chain, to total root respiration of both P. lanceolataand Z. mays, was not affected by light intensity during thedaytime. The alternative pathway was somewhat decreased at theend of the dark period, but not in the root tips (0–5mm) where it still contributed 56% in respiration. It was, therefore,concluded that photosynthesis is not a major factor in regulationof root growth in the species investigated. To see whether the effect of light intensity on root growthrate was via transpiration, plants of Z. mays were grown atdifferent air humidities. Both high humidity and low light intensityaffected the root morphology in such a way that the distancebetween the apex and the first laterals on the primary rootaxis increased. It is suggested that this effect on root morphologyis due to transpiration and the subsequent removal of root-producedinhibitors of lateral root growth; although light intensityalso affected the rate of dry matter accumulation of roots andthe rate was not affected by the humidity of the air. It is,therefore, concluded that the effect of light intensity on therate of dry matter accumulation of roots of Z. mays is not viaan effect on transpiration.