Effect of Irradiance on Chlorophyll Estimation with the Minolta SPAD-502 Leaf Chlorophyll Meter

Leaf chlorophyll content may be used as an indirect indicatorof crop nitrogen status. Chlorophyll meter values (SPAD values)taken with the Minolta SPAD-502 chlorophyll meter in the shadeplantOxalis acetosellaL. and in winter wheat (Triticum aestivumL.)varied by 15 and 8%, respectively, with variation in irradiance.The lowest SPAD-values were measured at high irradiance. Duringa natural night-day-night cycle SPAD values for winter wheatwere lowest in the middle of the day, highest at low irradianceat dusk and dawn and intermediate in darkness before dawn andafter dusk. The results indicate that irradiance during measurementshould be considered when using the Minolta SPAD-502 chlorophyllmeter for the estimation of crop N-status.Copyright 1998 Annalsof Botany Company Chlorophyll meter, nitrogen, irradiance,Oxalis acetosellaL.,Triticum aestivumL., winter wheat.     

Water Stress in Sunflower (Helianthus annuus L.) 2. Effects on Leaf Cells and Leaf Area

Three water stress treatments were applied at early, mid andlate stages of vegetative development in sunflower. The effectsof these stresses on leaf area, cell frequency, number of cellsper leaf and palisade cell area were examined. Final leaf area was reduced in all stress treatments. The largestreduction occurred in leaves that were unfolding or about tounfold at the commencement of stress. At full expansion of theseleaves cell frequency had increased, cell number had decreasedand cell area had decreased. These results help to explain theeffects of stress on final leaf area, especially the reducedarea of the lower leaves that are most sensitive to water stress. Helianthus annuus L., sunflower, water stress, node number, leaf area     

Leaf Growth in Cotton (Gossypium hirsutum L.) 1. Growth in Area of Main-stem and Sympodial Leaves

A model is presented for growth of individual and successivemain-stem leaves of cotton, based on a series of indoor experimentsand data sets from the literature. Three variable parametersare used to describe individual leaf growth: relative growthrate of meristematic tissue (R¹), relative rate of approachof final area (R₂) and a ‘position parameter’ (t_0.5)which governs the transition from meristematic to extensiongrowth. Final area of a leaf does not occur in the model asa deterministic quantity but it is a result of the processesduring growth. The model generates successive mainstem leavesand sympodial leaves as an integrated system. Assimilate shortagesoccurring in the plant operate on R₁ leading to the characteristicchange of final leaf area along the mainstem. Gossypium hirsutumL., cotton, leaf growth, relative growth rate, meristematic tissue, extension growth, mathematical model     

The Significance of Leaf Area in Evapotranspiration

Evapotranspiration rates increased with leaf area and fell withits decline at the time of flowering of cotton plants (Gossypiumbarbadense L.) and this relation was maintained while the rateof pan evaporation was steadily decreasing. A similar relationshipwas established for the hyacinth bean (Dolichos lablab L.) butevapotranspiration dropped at the onset of flowering, at thetime when leaf area was still increasing and the evaporatingpotential of the atmosphere began to rise. Evapotranspirationper unit area of leaf surface per day declined progressivelywith age for both species.     

Leaf area index determination in an alder forest: a comparison of three methods

Leaf area index (LAI) of a stand of adult black alder trees(Alnus glutinosa L., Gaertn.) was determined by means of threeindependent methods. (1) The seasonal course of LAI was directlyobtained by counting leaves in situ and adding up their areas,estimated from harvested subsamples of leaves. (2) The seasonalvariation of LAI in the stand was estimated using the Li-CorLAI-2000 PCA in parallel and with this instrument a VegetationArea Index (VAI, projected area of all phyto-elements) was actuallymeasured. (3) Maximum LAI was calculated from leaf litter collectionstaking into account specific leaf area within different layersof the alder crown. Direct LAI estimates (1) and calculationsfrom leaf litter (3) revealed the same figure of maximum LAI(4.8). This LAI was reached in August. The LAI-2000 PCA capturedthe seasonal variation and underestimated, by 11% on average,the LAI obtained directly. Compared with results gained withother broad-leaved tree species the LAI-2000 PCA values foralder were reliable. It is suggested that this is due to thehorizontal homogeneous structure of the main leaf layer. Thisis in the periphery of the crown, where 90% of the light interceptionoccurs. Taking the het-erogeneity into account a satisfactorycompatibility of the three methods applied to the alder standwas achieved. Key words: Alnus glutinosa, leaf area index, in situ counting, LAI-2000 PCA, litter collections     

Leaf Characteristics and Net Gas Exchange of Diploid and Autotetraploid Citrus

The effect of tetraploidy on leaf characteristics and net gasexchange was studied in diploid (2x ) and autotetraploid (4x) ‘Valencia’ sweet orange (Citrus sinensis (L.)Osb.) and ‘Femminello’ lemon (Citrus limon (L.)Burm. f.) leaves. Comparisons between ploidy levels were madeunder high irradiance (I) in a growth chamber or low total Iin a glasshouse. Tetraploids of both species had thicker leaves,larger mesophyll cell volume and lower light transmittance thandiploids regardless of growth I. Mesophyll surface area perunit leaf area of 2x leaves was 5–15% greater than on4x leaves. Leaf thickness and mesophyll cell volume were greaterin high I leaves than low I leaves. In high I, average leafarea was similar for 2x and 4x leaves, whereas in low I it was30% greater in 4x than in 2x leaves. Nitrogen and chlorophyllconcentration per cell increased with ploidy level in both growthconditions. The ratio of chlorophyll a:b was 25% greater in2x than in 4x leaves. When net CO₂assimilation rate (A_CO2) wasbased on leaf area, 4x orange leaves had 24–35% lowerA_CO2than their diploids. There were no significant differencesin A_CO2between 2x and 4x orange or lemon leaves when expressedon a per cell basis. Overall, lower A_CO2per unit leaf area oftetraploids was related to increase in leaf thickness, largermesophyll cell volume, the decrease in mesophyll area exposedto internal air spaces, and the lower ratio between cell surfaceto cell volume. Such changes probably increased the resistanceto CO₂diffusion to the site of carboyxlation in the chloroplasts. Cell volume; chlorophyll; irradiance; leaf thickness; nitrogen; photosynthesis; ploidy; Citrus limon ; C. sinensis ; ‘Valencia’ sweet orange; ‘Femminello’ lemon     

Growth Rate, Photosynthesis and Respiration in Relation to Leaf Area Index

This work examined three possible explanations of growth rateresponses to leaf area index (LAI) in which growth rate perunit of ground area (crop growth rate, CGR) increased to a plateaurather than decreasing above an optimum LAI at which all lightwas intercepted. Single leaf photosynthetic measurements, andwhole plant 24 h photosynthesis and respiration measurementswere made for isolated plants and plants in stands using Amaranlhushybridus, Chenopodium album, and two cultivars of Glycine maxgrown at 500 and 1000 µimol m^–2 S^–1 photosyntheticphoton flux density at 25 °C. CGR, relative growth rate(RGR), and LAI were determined from 24 h carbon dioxide exchangeand leaf area and biomass measurements. Respiration increasedrelative to photosynthesis with crowding in A. hybridus andthere was an optimum LAI for CGR. In contrast, the ratio ofrespiration to photosynthesis was constant across plant arrangementin the other species and they had a plateau response of CGRto LAI. Neither increased leaf photosynthetic capacity at highLAI nor a large change in biomass compared to the change inLAI could account for the plateau responses. It was calculatedthat maintenance respiration per unit of biomass decreased withdecreasing RGR in C. album and G. max, but not A. hybridus,and accounted for the plateau response of CGR to LAI. Sincesimilar decreases in maintenance respiration per biomass atlow RGR have been reported for several other species, a constantratio of respiration to photosynthesis may occur in more speciesthan constant maintenance respiration per unit of biomass. Amaranlhus hybridus L., Chenopodium album L., Glycine max L Merr, soybean, photosynthesis, respiration, growth, leaf area index     

Growth Rate and Water Relations of Citrus Leaf Flushes

Growth rates of seasonal leaf flushes of ‘Valencia’orange [Citrus sinensis (L.) Osbeck] were measured and waterrelations characteristics of young (new) and over-wintered (old)citrus leaves were compared. New flush leaves had lower specificleaf weights and lower midday leaf water potentials than comparablyexposed old leaves. Spring and summer flush new leaves had higherosmotic potentials than old leaves. These differences becamenon-significant as the new leaves matured. During summer conditions,water-stressed new leaves reached zero turgor and stomatal conductancealso began to decrease in them at higher leaf water potentialsthan in old leaves. Old leaves were capable of maintaining openstomata at lower leaf water potentials. Opened flowers and newflush leaves lost more water, on a dry weight basis, than flowerbuds, fruit or mature leaves. The results illustrate differencesin leaf water potential and stomatal conductance which can beattributed to the maintenance of leaf turgor by decreases inleaf osmotic potentials as leaves mature. These changes in citrusleaf water relations are especially important since water stressresulting from high water loss rates of new tissues could reduceflowering and fruit set. Citrus sinensis (L.) Osbeck, orange, Citrus paradisi Macf., grapefruit, growth rate, leaf water relations, osmotic potential, water potential, stomatal conductance     

Visible foliar injury caused by ozone alters the relationship between SPAD meter readings and chlorophyll concentrations in cutleaf coneflower

The ability of the SPAD-502 chlorophyll meter to quantify chlorophyll amounts in ozone-affected leaves of cutleaf coneflower (Rudbeckia laciniata var. digitata) was assessed in this study. When relatively uninjured leaves were measured (percent leaf area affected by stipple less than 6%), SPAD meter readings were linearly related to total chlorophyll with an adjusted R ² of 0.84. However, when leaves with foliar injury (characterized as a purple to brownish stipple on the upper leaf surface affecting more than 6% of the leaf area) were added, likelihood ratio tests showed that it was no longer possible to use the same equation to obtain chlorophyll estimations for both classes of leaves. Either an equation with a common slope or a common intercept was necessary. We suspect several factors are involved in altering the calibration of the SPAD meter for measuring chlorophyll amounts in visibly ozone-injured leaves, with the most likely being changes in either light absorption or scattering resulting from tissue necrosis.     

Comparison of Chlorophyll Meter Readings with Leaf Chlorophyll Concentration in Amaranthus vlitus: Correlation with Physiological Processes1

The portable chlorophyll meter (SPAD-502) has been successfully used for a rapid and direct estimation of total chlorophyll content (TCHL) in the leaves of some crops. In this work, SPAD-502 meter readings and TCHL concentration were compared for the leaves of Amaranthus vlitus L., a common weed. SPAD readings were linearly and positively correlated to TCHL concentration in the leaves. A linear correlation was also shown between SPAD-502 readings and some physiological parameters of the leaves, such as photosynthesis, transpiration, and stomatal conductance.     

Effects of Temperature and Leaf Position on Leaf Area Expansion of Kiwifruit (Actinidia deliciosa) Shoots: Development of a Modelling Framework

This paper describes mathematically the effects of temperatureand position on the expansion of leaves along a kiwifruit (Actinidiadeliciosa) shoot, taking into account shoot morphology. Theleaves were grouped into three zones along the shoot: initialcluster leaves (first zone); the rest of the leaves that werepreformed during the previous season (second zone); and leavesthat were initiated during the current season (third zone).After opening of the initial cluster, the leaves appeared atconstant rates for each of the two temperature treatments considered.The expansion of individual leaves was modelled by a growthfunction with the parameters: final area; duration of the growthwindow; centre of the growth window (timing of expansion); andlower asymptote. Within the first two zones, the pattern ofleaf expansion was affected by nodal position, with basal leaveshaving higher initial rates of expansion than distal leaves.The timing of expansion was linear with respect to the nodalposition within each of the zones, with the slope independentof temperature for the first zone. The slopes of the timingof expansion for the second and third zones depended on temperatureand were correlated for each temperature treatment. Final leafarea was a function of leaf position in the first zone and afunction of timing of leaf expansion for distal leaves startingfrom leaf 10. Temperature had no effect on final leaf area inthe first zone. For the rest of the leaves, temperature affectedfinal leaf area indirectly, through the timing of leaf expansion.The effect of temperature on the growth window of individualleaves within the first zone was less than that for the restof the leaves. However, simulated values for the total leafarea of shoots using the average shoot growth window showedgood agreement with experimental values.Copyright 2001 Annalsof Botany Company Actinidia deliciosa‘Hayward’, shoot development, individual leaf area, temperature effect, positional effects, modelling     

Methods of Estimating the Leaf Area of Grasses from Linear Measurements

This paper is concerned only with estimates of leaf area obtainedfrom linear measurements on the leaves. Past investigationson grasses and on other species are briefly reviewed. Variousweighted regressions are examined, the object being to obtaina satisfactory estimate of the constant k in the predictionequation A = kLB (L = length of leaf; B = breadth at a pointmidway along the length; A = area). Sets of data from four differentgrasses are examined. The formula A = 0?9O5LB is consistentwith all these data. The practical use of the technique is discussed.     

Profiles of Leaf Senescence During Reproductive Growth of Sunflower and Maize

We investigated the effect of reproductive growth on the profilesof leaf senescence in maize (Zea mays L.) and sunflower (Helianthusannuus L.). Leaf senescence after flowering was assessed usingboth structural (leaf chlorophyll, nitrogen and dry matter)and functional (photosynthesis) variables in undisturbed plants(+G) and in plants in which grain set was prevented (-G). Twoweeks after flowering, lack of grain accelerated senescencein maize and delayed senescence in sunflower as indicated byleaf chlorophyll; leaf nitrogen and dry matter were less sensitiveresponse variables. Lack of interaction between reproductivetreatment and leaf position indicates that the senescence signal,whatever its nature, was equally effective throughout the plantin both species. In both species, feedback inhibition of photosynthesiswas first detected 30–35 d after flowering; excess carbohydratein the leaves was therefore an unlikely trigger of acceleratedsenescence in maize. As reproductive development progressed,differences between +G and -G plants were more marked in sunflower,and tended to disappear or reverse in maize. In sunflower, interactionsbetween leaf position and reproductive treatment—attributableto the local effect of grain—were detected around 20–27d after flowering. Copyright 2000 Annals of Botany Company Helianthus annuus, Zea mays, chlorophyll, light, nitrogen, photosynthesis, reproductive growth, senescence, source-sink, SPAD.     

The Effect of Shade During Leaf Expansion on Photosynthesis by White Clover Leaves

In three experiments measurements of photosynthesis were madeon single leaves of white clover (Trifolium repens L.) on threecultivars grown in a controlled environment. Plants which had grown under an irradiance of 30 J m^–2s^–1, or in shade within a simulated mixed sward, producedleaves with photosynthetic capacities some 30 per cent lowerthan did plants grown at 120 J m^–2 s^–1 without shade.There were no differences between treatments either in photosynthesismeasured at 30 J m^–2 s^–1, or in respiration ratesper unit leaf dry weight. Respiration per unit leaf area washigher in the plants grown at 120 J m^–2 s^–1, reflectingthe lower specific leaf area of these leaves. There were nodifferences between the three cultivars examined. Leaves which were removed from the shade of a simulated swardshortly after becoming half expanded achieved photosyntheticcapacities as high as those which were in full light throughouttheir development. It is suggested that it is this characteristicwhich enables clover plants growing in an increasingly densemixed sward to produce a succession of leaves of high photosyntheticcapacity, even though each lamina only reaches the top of thesward at a relatively late stage in its development. Trifolium repens L., white clover, photosynthesis, leaf expansion, shade, specific leaf area, stomatal conductance     

The Dynamics of Leaf Growth and Photosynthetic Capacity in Capsicum frutescens L.

In Capsicum frutescens L. cv. California Wonder the specificleaf weight (dry weight per unit laminar area) at leaf unfoldingis three times higher in the eighth leaf than in the first leafproduced. Intermediate leaves exhibit a trend between the twoThe change in specific leaf weight during laminar expansionis greatest in leaf 1 and least (sometimes zero) in leaf 8.Large changes in specific leaf weight during laminar expansionare associated with a large degree of palisade cell expansion,while leaves showing smaller rates of change have less palisadecell expansion but cell division is more evident. At leaf unfoldingthe fraction I protein content per unit laminar area is higherin upper than in lower leaves. Ribulose diphosphate carboxylaseactivity per unit laminar area and ¹⁴CO₂ fixation per unit laminararea have a similar pattern of development in all leaves andshow no correlation with the changes in specific leaf weight.The peak of activity in all leaves occurs when the laminar areais 10 cm². These results are compared with previous data onlaminar expansion and are seen as in accord with current ideason leaf growth.     

Relationship between Leaf Structure and Gas Exchange in Wheat Leaves at Different Insertion Levels

Net photosynthesis rate (P_n), stomatal conductance to CO₂ andresidual conductance to CO₂ were measured in the last six leaves(the sixth or flag leaf and the preceding five leaves) of Triticumaestivum L. cv. Kolibri plants grown in Mediterranean conditions.Recently fully expanded leaves of well-watered plants were alwaysused. Measurements were made at saturating photosynthetic photonflux density, and at ambient CO₂ and O₂ levels. The specificleaf area, total organic nitrogen content, some anatomical characteristics,and other parameters, were measured on the same leaves usedfor gas exchange experiments. A progressive xeromorphic adaptation in the leaf structure wasobserved with increasing leaf insertion levels. Furthermore,mesophyll cell volume per unit leaf area (V_mes/A) decreasedby 52·6% from the first leaf to the flag leaf. Mesophyllcell area per unit leaf area also decreased, but only by 24·5%.However, nitrogen content per unit mesophyll cell volume increasedby 50·6% from the first leaf to the flag leaf. This increasecould be associated to an observed higher number of chloroplastcross-sections per mm² of mesophyll cell cross-sectional areain the flag leaf: values of 23000 in the first leaf and 48000in the flag leaf were obtained. P_n per unit leaf area remainedfairly constant at the different insertion levels: values of33·83±0·93 mg dm^–2 h^–1 and32·32±1·61 mg dm^–2 h^–1 wereobtained for the first leaf and the flag leaf, respectively.Residual conductance, however, decreased by 18·2% fromthe first leaf to the flag leaf. Stomatal conductance increasedby 41·7%. The steadiness in P_n per unit leaf area across the leaf insertionlevels could be mainly accounted for by an opposing effect betweena decrease in V_mes/A and a more closely packed arrangement ofphotosynthetic apparatus. Adaptative significance of structuralchanges with increasing leaf insertion levels and the steadinessin P_n per unit leaf area was studied. Key words: Photosynthesis, structure, wheat     

Short and Long Term Fluctuations of the Leaf Mass Per Area of Tomato Plants--Implications for Growth Models

The leaf mass per unit leaf area (LMA) is a key variable inmany growth models, since it is often used to predict leaf areaexpansion from leaf dry weight increase, orvice versa. Influencesof source-sink balance on leaf area, leaf dry weight, LMA, andleaf content in non-structural carbohydrates were investigatedin glasshouse tomato crops. The source-sink balance was manipulatedby artificial shading, CO₂enrichment or fruit removal usingdifferent tomato cultivars. Leaf area was hardly affected bycompetition for assimilates except under extreme conditions.In contrast, leaf dry weight, and consequently LMA, underwentlarge and rapid fluctuations in response to any factor thatchanged source and sink activities. A 60% reduction of photosyntheticallyactive radiation involved a 24% decrease in LMA after 10 d.Carbon dioxide enrichment and fruit removal induced about a45% and 15% increase in LMA, respectively, on plants with twofruiting trusses, but hardly affected LMA of producing plants.No significant cultivar effect could be identified. Changesin starch and soluble sugar content in leaves accounted foronly 29% of diurnal variations in LMA, suggesting regular fluctuationsof other components. We propose that structural LMA varies betweena maximum and a minimum value according to the ratio of assimilatesupply and demand during leaf development. Leaf area is independentof the supply of assimilates when the minimum structural LMAis realised. When the maximum structural LMA is attained, astorage pool of assimilates may accumulate in leaves duringperiods of high supply and low demand. We present a model includingthese hypotheses, which predicts structural and non-structuralLMA variations of plants with different source-sink ratios.Copyright1998 Annals of Botany Company Tomato,Lycopersicon esculentumMill., SLA, SLW, leaf growth, vegetative sink strength, assimilate competition, source-sink ratio, non-structural carbohydrate, models.     

Macroscopic Features of Wind Damage to Leaves of Acer pseudoplatanus L. and Its Relationship with Season, Leaf Age, and Windspeed

Exposure to wind produces permanent abrasive damage to Acerpseudoplatanus L. leaves in the form of brown lesions, holes,distortions and tearing of the lamina. The zones of leaf thatare readily damaged change as the leaves expand and their topographyis altered. In the field, most damage was sustained early in the seasondespite the persistence of wind throughout the year. Wind tunnelstudies demonstrated that young expanding leaves were far moresusceptible to wind than mature leaves and that the percentagearea damaged increased linearly with windspeed. Acer pseudoplatanus L., sycamore, wind, leaf damage     

Leaf and Tiller Production of Prairie Grass (Bromus willdenowii Kunth) and Two Ryegrass (Lolium) Species

A detailed morphological study of three prairie grass cultivars(Bromus willdenowii Kunth) was conducted under ‘vegetative’and ‘reproductive’ growth conditions (short andlong photoperiods) and at different temperatures. Perennialryegrass (Lolium perenne L.) and Westerwolds ryegrass (Loliummuhiflorum Lam.) were compared during vegetative growth. Prairie grass had higher leaf appearance rates (leaves per tillerper day) and lower site filling (tillers per tiller per leafappearance interval) than the ryegrass species. Tillering rates(tillers per tiller per day) were also lower, except under vegetativeconditions at 4C. Low tiller number in prairie grass was notdue to lack of tiller sites but a result of poor filling ofthese sites. Lower site filling occurred because of increaseddelays in appearance of the youngest axillary tiller and lackof axillary tillers emerging from basal tiller buds. In prairiegrass, no tillers came from coleoptile buds while only occasionallydid prophyll buds develop tillers. Low tiller number in prairiegrass was compensated for by greater tiller weight. Prairiegrass had more live leaves per tiller, greater area per leafand a high leaf area per plant. Considerable variation between cultivars was found in prairiegrass. The cultivar ‘Bellegarde’ had high leaf appearance,large leaves and rapid reproductive development, but had lowlevels of site filling, tillering rate, final tiller numberand herbage quality during reproductive growth. ‘Primabel’tended to have the opposite levels for these parameters, while‘Grasslands Matua’ was intermediate and possiblyprovided the best balance of all plant parameters. prairie grass, Bromus willdenowii Kunth, perennial ryegrass, Lolium perenne L., Westerwolds ryegrass, Lolium multiflorum Lam., temperature, photoperiod, leaf appearance, leaf area, tillering, site filling, tillering sites, yield     

Leaf Primordium Initiation and Expanded Leaf Production are Co-ordinated through Similar Response to Air Temperature in Pea (Pisum sativumL.)

Accurate prediction of the timing of leaf area development isessential to analyse and predict the responses of crops to theenvironment. In this paper, we analyse the two processes determiningthe chronology of leaf development—initiation of leafprimordia by the shoot meristem and production of expanded leavesout of the shoot tip—in several pea (Pisum sativumL.)cultivars in response to air temperature and plant growth rate.Contrasting levels of air temperature and plant growth rateduring leaf development were induced by a wide range of sowingdates and plant densities in glasshouse or field experiments.Full leaf expansion was found to occur one phyllochron afterfull leaf unfolding, whatever the leaf nodal position. Primordiuminitiation and expanded leaf production rates presented similarquantitative responses to air temperature (linear response andcommonx-intercept), whatever the plant growth rate, cultivaror period of cycle. As a consequence, they were co-ordinatedand the numbers of initiated primordia or expanded leaves wereeasily deduced from simple visual observation of leaf unfolding.The change, over time, of the numbers of initiated leaf primordiaand fully expanded leaves correlated with cumulated degree-days,with stable relationships in a wide range of environmental conditions.Two phases, with different production rates, had to be considered.These results allowed us to predict accurately the beginningand the end of individual leaf development from daily mean airtemperatures. The relationships obtained here provide an effectiveway of analysing and predicting leaf development responses tothe environment. Pisum sativumL.; pea; number of leaf primordia; number of leaves; temperature; modelling