Baobab trees (Adansonia) in Madagascar use stored water to flush new leaves but not to support stomatal opening before the rainy season

Baobab trees (Adansonia, Bombacaceae) are widely thought to store water in their stems for use when water availability is low. We tested this hypothesis by assessing the role of stored water during the dry season in three baobab species in Madagascar. In the dry season, leaves are present only during and after leaf flush. We quantified the relative contributions of stem and soil water during this period through measures of stem water content, sap flow and stomatal conductance. Rates of sap flow at the base of the trunk were near zero, indicating that leaf flushing was almost entirely dependent on stem water. Stem water content declined by up to 12% during this period, yet stomatal conductance and branch sap flow rates remained very low. Stem water reserves were used to support new leaf growth and cuticular transpiration, but not to support stomatal opening before the rainy season. Stomatal opening coincided with the onset of sap flow at the base of the trunk and occurred only after significant rainfall.     

Experimental Studies of the Factors Controlling Transpiration: I. APPARATUS AND EXPERIMENTAL TECHNIQUE

Apparatus and experimental techniques are discussed for usein the investigation of transpiration rate of leaves as controlledby stomatal aperture and leaf water content. The leaf chambers used and the methods adopted for the estimationof the water transpired are described. The designs of the porometer cups used for the different typesof leaves (Pelargonium and wheat) employed in the work are described.To obviate the difficulty that stomata within the cup behaveabnormally, the design employed makes possible a removal ofthe cup from the leaf except during the short periods requiredto estimate stomatal resistance to air flow at intervals duringthe course of an experiment. In these experiments the water content of the leaf is changedat will by interrupting the water-supply and re-establishingit to permit recovery from wilting. The methods used to carryout this cycle of operations are fully dealt with. Determinationsof the transpiration and absorption rates during the experimentand of the final leaf water content make it possible to followchanges in leaf water content throughout the experiment. An account is given of the methods used for varying the speedof flow, the humidity, and the CO₂ concentration of the airstreams.     

Direct and indirect effects of elevated CO2 on transpiration from Quercus myrtifolia in a scrub-oak ecosystem

Elevated atmospheric carbon dioxide (C_a) usually reduces stomatal conductance, but the effects on plant transpiration in the field are not well understood. Using constant‐power sap flow gauges, we measured transpiration from Quercus myrtifolia Willd., the dominant species of the Florida scrub‐oak ecosystem, which had been exposed in situ to elevated C_a (350 µmol mol ^{? 1} above ambient) in open‐top chambers since May 1996. Elevated C_a reduced average transpiration per unit leaf area by 37%, 48% and 49% in March, May and October 2000, respectively. Temporarily reversing the C_a treatments showed that at least part of the reduction in transpiration was an immediate, reversible response to elevated C_a. However, there was also an apparent indirect effect of C_a on transpiration: when transpiration in all plants was measured under common C_a, transpiration in elevated C_a‐grown plants was lower than that in plants grown in normal ambient C_a. Results from measurements of stomatal conductance (g_s), leaf area index (LAI), canopy light interception and correlation between light and g_s indicated that the direct, reversible C_a effect on transpiration was due to changes in g_s caused by C_a, and the indirect effect was caused mainly by greater self‐shading resulting from enhanced LAI, not from stomatal acclimation. By reducing light penetration through the canopy, the enhanced self‐shading at elevated C_a decreased stomatal conductance and transpiration of leaves at the middle and bottom of canopy. This self‐shading mechanism is likely to be important in ecosystems where LAI increases in response to elevated C_a.     

The Effect of Reduced Hydraulic Conductance on Stomatal Conductance and Xylem Cavitation

The vulnerability of xylem conduits to cavitation theoreticallydetermines the maximum flow rate of water through plants, andhence maximum transpiration (E), stomatal conductance (g_s),and leaf area (A₁. Field-grown Betula occidentalis with a favourablewater supply exhibit midday xylem pressures (     

Water relations and hydraulic control of stomatal behaviour in bell pepper plant in partial soil drying

Two experiments, a split-root experiment and a root pressurizing experiment, were performed to test whether hydraulic signalling of soil drying plays a dominant role in controlling stomatal closure in herbaceous bell pepper plants. In the split-root experiment, when both root parts were dried, synchronous decreases in stomatal conductance (g_s), leaf water potential (LWP) and stem sap flow (SF_stem) were observed. The value of g_s was found to be closely related to soil water potential (SWP) in both compartments. Tight relationships were observed between g_s and stem sap flow under all conditions of water stress, indicating a complete stomatal adjustment of transpiration. When the half-root system has been dried to the extent that its water uptake dropped to almost zero, declines in g_s of less than 20% were observed without obvious changes in LWP. The reduced plant hydraulic conductance resulting from decreased sap flow and unchanged LWP may be a hydraulic signal controlling stomatal closure; the results of root pressurizing supported this hypothesis. Both LWP and g_s in water-stressed plants recovered completely within 25 min of the application of root pressurizing, and decreased significantly within 40 min after pressure release, indicating the hydraulic control of stomatal closure. Our results are in contrast to those of other studies on other herbaceous species, which suggested that chemical messengers from the roots bring about stomatal closure when plants are in water stress.     

Stomatal Regulation by Aminoacetonitrile, a Photor espiration Inhibitor

The effect of aminoacetonitrile (AAN), a specific inhibitorof photorespiration, on photosynthesis and transpiration ofrice and maize leaves, C₃ and C₄ plants, respectively, was investigated.Application of AAN to the rice leaf in atmospheric air, thatis, under a photorespiratory condition, reduced both photosynthesisand transpiration, whereas its application to rice leaf in 2%O₂ air, that is, under a nonphotorespiratory condition, didnot affect photosynthesis or transpiration. Application of AANto the maize leaf did not affect photosynthesis or transpiration.Theseresults suggest that stomatal behavior is closely linked tophotorespiration. (Received March 12, 1987; Accepted August 21, 1987)     

Stomatal responses to light and CO2 in greening wheat leaves

The development of two types of stomatal transpiration, oneinduced by light (light-induced stomatal transpiration) andthe other induced by CO₂-free air in the dark (CO₂-sensitivestomatal transpiration), in greening leaves of wheat (Triticumaestivum L.) was studied in respect to the development of CO₂uptake and chlorophyll formation. Light-induced stomatal transpirationwas not observed at all in etiolated leaves and was generatedafter 3 hr of illumination for greening, when the activity ofCO₂ uptake was generated. CO₂-sensitive stomatal transpirationwas low in etiolated leaves and started to increase at the sametime during greening as the start of CO₂ uptake. The activitiesof both light-induced and CO₂-sensitive stomatal transpirationincreased as the activity of CO₂ uptake and the chlorophyllcontent increased. Pre-illumination of etiolated leaves for1 min followed by 4 hr of dark incubation eliminated the lagfor the development of the two types of stomatal transpirationand CO₂ uptake. (Received September 4, 1978; )     

Relative rates of delivery of xylem solute to shoot tissues: Possible relationship to sequential leaf senescence

The rates of delivery of regulatory solutes such as cytokinins and mineral ions from the roots to competing shoot tissues can influence rates of metabolism and development. A 15 min pulse of a synthetic xylem mobile and phloem-immobile solute, acid fuchsin, was used to quantify relative rates of solute delivery to competing organs on excised transpiring bean shoots (Phaseolus vulgaris L. cv. Contender) at different stages of development. Stem, flower and fruit tissues received comparatively low rates of solute delivery. The relative rate of solute delivery to newly opened leaves was initially low, but increased during rapid leaf expansion and then declined progressively as the leaves exceeded 70% of their final area. The relative rate of solute delivery to tissues of the basal primary leaves declined progressively from 2 weeks onwards. This decline appeared to be caused by progressive internally regulated increases in both stomatal resistances and hydraulic resistances to xylem flow up to and into the leaf blade. Thus combined abaxial and adaxial stomatal resistance values in the primary leaves (Rs) increased from 3 to ≥ 7 s cm^?1 between 2 and 5 weeks. Similarly, mean values for the connection resistances (Rc) to hydraulic flow into the primary leaves rose from 7 to 13 TPa · s · m^?1 between 2 and 4 weeks. In the same period pathway resistance from stem to primary leaf petioles (Rp), as determined by direct pressure flow assay, increased from 7 to 15 TPa · s · m^?1. Senescence-associated declines in protein and chlorophyll levels in the primary leaves were initiated in parallel with, or after, declines in relative rates of solute delivery. The provision of extra illumination at the basal leaf level between 2 and 5 weeks did not prevent declines in chlorophyll and soluble protein or increases in stomatal resistance. We suggest that internally programmed changes in the hydraulic architecture of the plant progressively divert xylem-transported root supplies of nutrients and cytokinins from basal to more apical leaves and thus regulate the progressive senescence of leaves from the base upwards.     

Field Studies of Cereal Leaf Growth: II. THE RELATION BETWEEN AUXANOMETER AND DISSECTION MEASUREMENTS OF LEAF EXTENSION AND THEIR RELATION TO CROP LEAF AREA EXPANSION

Destructive measurements of laminar, sheath, and internode lengthmade on spring barley and winter wheat enabled the contributionof these phytomer components to apparent leaf extension rate(R_E), as measured by auxanometer or rule, to be determined.R_E for early formed leaves consisted entirely of laminar andsheath extension. However, during stem extension, internodeextension also contributed to measured R_E. For the last twoleaves of winter wheat, the extension of internode below thephytomer being measured contributed to R_E. For these leaves,R_E should therefore be interpreted cautiously. Calculating mainstem and tiller leaf areas, and allowing for changes of leafwidth with leaf number, showed that leaf extension was closelyrelated to crop leaf area expansion.     

Changing Responses of Stomata to Abscisic Acid and CO2 as Leaves and Plants Age

Willmer, C. M., Wilson, A. B. and Jones, H. G. 1988. Changingresponses of stomata to abscisic acid and CO₂ as leaves andplants age.—J. exp. Bot. 39: 401–410. Stomatal conductances were measured in ageing leaves of Commelinacommunis L. as plants developed; stomatal responses to CO₂ andabscisic acid (ABA) in epidermal strips of C. communis takenfrom ageing leaves of developing plants and in epidermal stripsfrom the same-aged leaves (the first fully-expanded leaf) ofdeveloping plants were also monitored. Stomatal behaviour wascorrelated with parallel measurements of photosynthesis andleaf ABA concentrations. Stomatal conductance in intact leavesdecreased from a maximum of 0-9 cm s^{– 1} at full leaf expansionto zero about 30 d later when leaves were very senescent. Conductancesdeclined more slowly with age in unshaded leaves. Photosynthesisof leaf slices also declined with age from a maximum at fullleaf expansion until about 30 d later when no O₂ exchange wasdetectable. Exogenously applied ABA (0.1 mol m^{– 3}) didnot affect respiration or photosynthesis. In epidermal stripstaken from ageing leaves the widest stomatal apertures occurredabout 10 d after full leaf expansion (just before floweringbegan) and then decreased with age; this decrease was less dramaticin unshaded leaves. The inhibitory effects of ABA on stomatalopening in epidermal strips decreased as leaves aged and wasgreater in the presence of CO₂ than in its absence. When leaveswere almost fully-senescent stomata were still able to open.At this stage, guard cells remained healthy-looking with greenchloroplasts while mesophyll cells were senescing and theirchloroplasts were yellow. Similar data were obtained for stomatain epidermal strips taken from the same-aged leaves of ageingplants. The inhibitory effects of ABA on stomatal opening alsodecreased with plant age. In ageing leaves both free and conjugated ABA concentrationsremained low before increasing dramatically about 30 d afterfull leaf expansion when senescence was well advanced. Concentrationsof free and conjugated ABA remained similar to each other atall times. It is concluded that the restriction of stomatal movements inintact leaves as the leaves and plants age is due mainly toa fall in photosynthetic capacity of the leaves which affectsintracellular CO₂ levels rather than to an inherent inabilityof the stomata to function normally. Since stomatal aperturein epidermal strips declines with plant and leaf age and stomatabecome less responsive to ABA (while endogenous leaf ABA levelsremain fairly constant until leaf senescence) it is suggestedthat some signal, other than ABA, is transmitted from the leafor other parts of the plant to the stomata and influences theirbehaviour. Key words: Abscisic acid, CO₂, Commelina, leaf age, senescence, stomatal sensitivity     

基于树干液流测定值进行尺度扩展的马占相思林段蒸腾和冠层气孔导度

应用Granier热消散探针,长期监测华南丘陵地马占相思(Acacia mangium)林14棵样树的树干液流(Sap flow),由此计算整树和林段的蒸腾速率,结合同步记录的环境因子,求算冠层平均气孔导度(G_c)。Granier探针的灵敏度较高,能精确测定即使是微弱的液流活动。观测结果显示,树木个体之间的液流密度(J_s)和整树蒸腾(E_t)受树形特征影响较大。马占相思林径级大的树木个体数较少,但占据林段边材总面积和林段蒸腾的比例较大。J_s和E_t的日变化主要受光合有效辐射(Q_o)和空气水蒸气压亏缺(D)的控制,土壤含水量(θ)对较大胸径树木E_t的影响大于胸径较小的树木,个体之间J_s和E_t的差异随θ的下降而缩小。一年中,林段蒸腾(E)在光照和水热条件较好的7月最高,9~12月,由于土壤水分供应的减少致使E值降低,E对D的敏感性下降。G_c与主要环境因子的关系与E相似,如果θ长期偏低,G_c会明显下降,是造成E降低的主要原因。成熟马占相思林在光照充足、水热条件较好的情况下的蒸腾活动旺盛,但对土壤水分胁迫的忍受力较低。     

Genotypic variation in transpiration of coppiced poplar during the third rotation of a short‐rotation bio‐energy culture

The productivity of short‐rotation coppice (SRC) plantations with poplar (Populus spp.) strongly depends on soil water availability, which limits the future development of its cultivation, and makes the study of the transpirational water loss particularly timely under the ongoing climate change (more frequent drought and floods). This study assesses the transpiration at different scales (leaf, tree and stand) of four poplar genotypes belonging to different species and from a different genetic background grown under an SRC regime. Measurements were performed for an entire growing season during the third year of the third rotation in a commercial scale multigenotype SRC plantation in Flanders (Belgium). Measurements at leaf level were performed on specific days with a contrasted evaporative demand, temperature and incoming shortwave radiation and included stomatal conductance, stem and leaf water potential. Leaf transpiration and leaf hydraulic conductance were obtained from these measurements. To determine the transpiration at the tree level, single‐stem sap flow using the stem heat balance (SHB) method and daily stem diameter variations were measured during the entire growing season. Sap flow‐based canopy transpiration (E_c), seasonal dry biomass yield, and water use efficiency (WUE; g aboveground dry matter/kg water transpired) of the four poplar genotypes were also calculated. The genotypes had contrasting physiological responses to environmental drivers and to soil conditions. Sap flow was tightly linked to the phenological stage of the trees and to the environmental variables (photosynthetically active radiation and vapor pressure deficit). The total E_c for the 2016 growing season was of 334, 350, 483 and 618 mm for the four poplar genotypes, Bakan, Koster, Oudenberg and Grimminge, respectively. The differences in physiological traits and in transpiration of the four genotypes resulted in different responses of WUE.     

Response of citrus trees to modified radiation regime in semi-arid conditions

Citrus trees are characterized by a large canopy and low hydraulicconductivity. In Israel's semi-arid summer climate this couldcause transpiration to exceed water uptake and cause temporaryexcessive water deficits. It was hypothesized that reductionof radiative load would reduce transpiration and thus reducedeficits. Net radiation of lemon trees in the hottest season was reducedby shading hedgerows with reflective nets for approximatelyone month in both 1994 and 1995. Stem sap flow and climate variableswere measured continuously. Daily courses of leaf conductanceand leaf water potentials were measured on selected days. Midday net radiation below the dense and sparse shade net treatmentswas 47% and 73% of that above the control trees. Midday ‘sunlit’leaf temperatures below the nets were reduced by 2.7 and 1.6C,respectively. The reduction in net radiation caused large changes in leafconductance. Average midday sunlit leaf conductance measuredin 1995 under the dense and sparse treatments and control were4.1, 2.9 and 1.8mm s^–1, respectively (significantly differentat P <0.01). Similar differences in sunlit leaf conductancewere found in 1994. Shade leaf conductance was not affectedby the treatments. Daily total and midday sap flow under the dense net were reducedby 6–7% and 10–11%, respectively. Sap flow underthe sparse net did not change significantly in 1994, but in1995 daily and midday sap flows were reduced by 6% and 7%, respectively.Midday leaf water potentials increased by 0.2 and 0.1 MPa underdense shade in 1994 and 1995, respectively. Under sparse shademidday leaf water potentials increased by 0.1 MPa in 1994, butdid not change significantly in 1995. A modified Penman-Monteith model evaluated transpiration ifleaf conductance were constant in the different radiation environments.At leaf conductance levels found in the unshaded trees, denseshade was estimated to cause a 25% reduction in transpiration,while leaf conductance values found in trees under the denseshade would lead to an increase in transpiration of more than35% in unshaded trees. The ability of the tree to maintain almost constant transpirationin different radiation environments and thus avoid water deficitby adjusting the conductance of sunlit leaves is discussed interms of environmental influences and significance to the plant'swater balance. Key words: Tree transpiration, stomatal closure, climate modification, citrus     

Rapid hydraulic recovery in Eucalyptus pauciflora after drought: linkages between stem hydraulics and leaf gas exchange

In woody plants, photosynthetic capacity is closely linked to rates at which the plant hydraulic system can supply water to the leaf surface. Drought‐induced embolism can cause sharp declines in xylem hydraulic conductivity that coincide with stomatal closure and reduced photosynthesis. Recovery of photosynthetic capacity after drought is dependent on restored xylem function, although few data exist to elucidate this coordination. We examined the dynamics of leaf gas exchange and xylem function in Eucalyptus pauciflora seedlings exposed to a cycle of severe water stress and recovery after re‐watering. Stomatal closure and leaf turgor loss occurred at water potentials that delayed the extensive spread of embolism through the stem xylem. Stem hydraulic conductance recovered to control levels within 6 h after re‐watering despite a severe drought treatment, suggesting an active mechanism embolism repair. However, stomatal conductance did not recover after 10 d of re‐watering, effecting tighter control of transpiration post drought. The dynamics of recovery suggest that a combination of hydraulic and non‐hydraulic factors influenced stomatal behaviour post drought.     

Transpiration in response to variation in microclimate and soil moisture in southeastern deciduous forests

Responses of forests to changes in environmental conditions reflect the integrated behavior of their constituent species. We investigated sap flux-scaled transpiration responses of two species prevalent in upland eastern hardwood forests, Quercus alba in the upper canopy and Acer rubrum in the low to mid canopy, to changes in photosynthetically active radiation above the canopy (Q_o), vapor pressure deficit within the canopy (D), and soil moisture depletion during an entire growing season. Water loss before bud break (presumably through the bark) increased linearly with D, reaching 8% of daily stand transpiration (E_C) as measured when leaf area index was at maximum, and accounting for 5% of annual water loss. After leaves were completely expanded and when soil moisture was high, sap flux-scaled daily E_C increased linearly with the daily sum of Q_o. Species differences in this response were observed. Q. alba reached a maximum transpiration at low Q_o, while A. rubrum showed increasing transpiration with Q_o at all light levels. Daily E_C increased in response to daily average D, with an asymptotic response due to the behavior of Q. alba. Transpiration of A. rubrum showed a greater response to soil moisture depletion than did that of Q. alba. When evaluated at a half-hourly scale under high Q_o, mean canopy stomatal conductance (G_S) of individuals decreased with D. The sensitivity of G_S to D was greater in species with higher intrinsic G_S. Regardless of position in the canopy, diffuse-porous species in this and an additional, more mesic stand showed higher G_S and greater stomatal sensitivity to environmental variation than do ring-porous species.     

Regulation of water flux through trunks, branches, and leaves in trees of a lowland tropical forest

We studied regulation of whole-tree water use in individuals of five diverse canopy tree species growing in a Panamanian seasonal forest. A construction crane equipped with a gondola was used to access the upper crowns and points along the branches and trunks of the study trees for making concurrent measurements of sap flow at the whole-tree and branch levels, and vapor phase conductances and water status at the leaf level. These measurements were integrated to assess physiological regulation of water use from the whole-tree to the single-leaf scale. Whole-tree water use ranged from 379 kg day⁻¹ in a 35 m-tall Anacardium excelsum tree to 46 kg day⁻¹ in an 18 m-tall Cecropia longipes tree. The dependence of whole-tree and branch sap velocity and sap flow on sapwood area was essentially identical in the five trees studied. However, large differences in transpiration per unit leaf area (E) among individuals and among branches on the same individual were observed. These differences were substantially reduced when E was normalized by the corresponding branch leaf area:sapwood area ratio (LA/SA). Variation in stomatal conductance (g _s) and crown conductance (g _c), a total vapor phase conductance that includes stomatal and boundary layer components, was closely associated with variation in the leaf area-specific total hydraulic conductance of the soil/leaf pathway (G _t). Vapor phase conductance in all five trees responded similarly to variation in G _t. Large diurnal variations in G _t were associated with diurnal variation in exchange of water between the transpiration stream and internal stem storage compartments. Differences in stomatal regulation of transpiration on a leaf area basis appeared to be governed largely by tree size and hydraulic architectural features rather than physiological differences in the responsiveness of stomata. We suggest that reliance on measurements gathered at a single scale or inadequate range of scale may result in misleading conclusions concerning physiological differences in regulation of transpiration. Received: 1 October 1997 / Accepted: 6 March 1998     

Does foliage on the same branch compete for the same water? Experiments on Douglas-fir trees

Do branchlets within a branch have autonomous water supplies, or do they share a common water supply system? We hypothesized that if branchlets shared a common water supply, then stomatal conductance (g_s) on sunlit foliage would increase with reduced transpiration of competing foliage on the branch. We reduced transpiration of other foliage on the branch through bagging and shading, and we monitored the gas-exchange responses of the remaining sunlit foliage on the branch relative to control branches for several age classes of Douglas-fir trees (aged ~10 years, 20 years, and 450 years old). Contrary to our hypothesis, we found no increases in g_s in either young or old trees following transient reductions in the amount of transpiring leaf area. The diurnal change in water potential, mid-day stomatal closure and associated photosynthetic decline occurred at the same time and were of the same magnitude on both treated and untreated branches, with the exception of photosynthesis in one 450-year-old tree. Hydraulic conductance measurements of branch junctions indicate that xylem within branches is only partially interconnected which would reduce the effectiveness of shading as a means of increasing water supply to the remaining sunlit foliage. The lack of a response implies that when a branch is in partial shade, the remaining sunlit foliage has no advantage with respect to water status over foliage on a branch completely in the sun.     

Patterns of Stomatal Behaviour, Transpiration, and CO2 Exchange in Pea Following Infection by Powdery Mildew (Erysiphe pisi)

A comparison was made of stomatal behaviour, and related phenomena,between leaves of garden pea (Pisum sativum cv. Feltham First)inoculated with powdery mildew fungus (Erysiphe pisi) and uninfectedleaves on healthy plants. Twenty four hours after inoculation,stomata opened more widely in the light in infected leaves thanin healthy leaves. Thereafter, stomatal opening was progressivelyreduced by infection and stomata failed to close completelyin the dark until, 7 d after inoculation, all movements ceasedand stomata remained partly open. Transpiration in the lightfollowed closely the pattem of stomatal opening and, after anearly increase compared with healthy controls, was progressivelyreduced by infection. Evidence is presented that transpirationfrom the fungus was less than the reduction in transpiraationfrom the leaf which was caused when development of the myceliumincreased the boundary layer resistance of the leaf. Seven daysafter inoculation, transpiration in the dark was greater frominfected leaves than from healthy leaves because of partly openstomata in the dark. Net photosynthesis in infected leaves was reduced within 24h of inoculation to a level below that found in healthy leavesand thereafter it declined progressively. The initial reductionwas due to a transient increase in photorespiration, for whenthe glycolate pathway was inhibited by a 2% O₂ concentrationthere was no difference between the (gross) photosynthetic ratesof healthy and infected leaves. Changes in photorespirationrate were confirmed from the interpretation of the CO₂ burston darkening. Reduced stomatal opening was a contributory causeof the reduction in net photosynthesis in the later stages ofinfection. Since the rate of gross photosynthesis, but not therate of photorespiration, of infected plants fell below thatof healthy plants, and infected plants had a higher rate ofrelease of CO₂ in the dark than healthy plants from the thirdday after inoculation onwards, infected plants consume an increasinglygreater proportion of their photosynthate in respiratory processesthan do healthy plants. The CO₂ compensation point of infectedplants increased at every time of sampling after inoculation.     

Effects of scale insect herbivory and shading on net gas exchange and growth of a subtropical tree species (Guaiacum sanctum L.)

Summary The scale insect, Toumeyella sp., feeds exclusively on the subtropical hammock tree lignum vitae (Guaiacum sanctum L.). The combined effects of scale herbivory and shading on leaf gas exchange characteristics and growth of lignum vitae trees were studied using a factorial design. Trees grown in full sun or in 75% shade were manually infested with scale or left noninfested. Beginning 4 weeks after infestation, net CO₂ assimilation, stomatal conductance, transpiration, internal partial pressure of CO₂, and water-use efficiency were determined on single-leaves at 4-week intervals for trees in each treatment. At the end of the experiment, net CO₂ assimilation was determined for whole plants. Total leaf area, leaf, stem, and root dry weights, and leaf chlorophyll and nitrogen concentrations were also determined. Scale infested trees generally had lower net CO₂ assimilation, stomatal conductance, and transpiration rates as well as less leaf area, and root, stem, and leaf dry weights than noninfested trees. Twenty four weeks after the shade treatment was imposed, sun-grown trees had approximately twice the leaf area of shade-grown trees. Shade-grown trees compensated for the reduced leaf area by increasing their photosynthetic efficiency. This resulted in no difference in light saturated net CO₂ assimilation on a whole plant basis between sun-grown and shade-grown trees. Chlorophyll and nitrogen concentrations per unit leaf area were greater in leaves of shade-grown trees than in leaves of sun-grown trees. Shading and herbivory by Toumeyella sp. each resulted in decreased growth of Guaiacum sanctum. Scale insect herbivory did not result in greater detrimental effects on leaf gas exchange characteristics for shade-grown than for sun-grown trees. Herbivory by Toumeyella resulted in a greater decrease in tree growth for sun-grown than for shade-grown trees.     

Stomatal Responses of Variegated Leaves to CO2 Enrichment

The responses of stomatal density and stomatal index of fivespecies of ornamental plants with variegated leaves grown attwo mole fractions of atmospheric CO₂ (350 and 700 µmolmol^-1) were measured. The use of variegated leaves allowed anypotential effects of mesophyll photosynthetic capacity to beuncoupled from the responses of stomatal density to changesin atmospheric CO₂ concentration. There was a decrease in stomataldensity and stomatal index with CO₂ enrichment on both white(unpigmented) and green (pigmented) leaf areas. A similar responseof stomatal density and index was also observed on areas ofleaves with pigmentation other than green indicating that anydifferences in metabolic processes associated with colouredleaves are not influencing the responses of stomatal densityto CO₂ concentrations. Therefore the carboxylation capacityof mesophyll tissue has no direct influence on stomatal densityand index responses as suggested previously (Friend and Woodward1990 Advances in Ecological Research 20: 59-124), instead theresponses were related to leaf structure. The stomatal characteristics(density and index) of homobaric variegated leaves showed agreater sensitivity to CO₂ on green portions, whereas heterobaricleaves showed a greater sensitivity on white areas. These resultsprovide evidence that leaf structure may play an important rolein determining the magnitude of stomatal density and index responsesto CO₂ concentrations.Copyright 1995, 1999 Academic Press Leaf structure, photosynthesis, stomatal conductance, CO₂, stomatal density, stomatal index