Xylem cavitation and hydraulic control of stomatal conductance in Laurel (Laurus nobilis L.)

The possible link between stomatal conductance (g_L), leaf water potential ( Ψ _L) and xylem cavitation was studied in leaves and shoots of detached branches as well as of whole plants of Laurus nobilis L. (Laurel). Shoot cavitation induced complete stomatal closure in air‐dehydrated detached branches in less than 10 min. By contrast, a fine regulation of g_L in whole plants was the consequence of Ψ _L reaching the cavitation threshold ( Ψ _CAV) for shoots. A pulse of xylem cavitation in the shoots was paralleled by a decrease in g_L of about 50%, while Ψ _L stabilized at values preventing further xylem cavitation. In these experiments, no root signals were likely to be sent to the leaves from the roots in response to soil dryness because branches were either detached or whole plants were growing in constantly wet soil. The stomatal response to increasing evaporative demand appeared therefore to be the result of hydraulic signals generated during shoot cavitation. A negative feedback link is proposed between g_L and Ψ _CAV rather than with Ψ _L itself.     

Hydraulic architecture of leaf blades: where is the main resistance?

The hydraulic architecture of Laurus nobilis L. and Juglans regia L. leaves was studied using three different approaches: (1) hydraulic measurements of both intact leaves and of leaves subjected to treatments aimed at removing the extra‐vascular resistance; (2) direct measurements of the vascular pressure with a pressure probe; and (3) modelling the hydraulic architecture of leaf venation system on the basis of measurements of vein densities and conductivities. The hydraulic resistance of leaves (R_leaf) either cut, boiled or frozen–thawed was reduced by about 60 and 85% with respect to control leaves for laurel and walnut, respectively. Direct pressure drop measurements suggested that 88% of the resistance resided outside the vascular system in walnut. Model simulations were in agreement with these results provided vein hydraulic conductance was 0.12–0.28 that of the conductance predicted by Poiseuille's law. The results suggest that R_leaf is dominated by substantial extra‐vascular resistances and therefore contrast with the conclusions of recent studies dealing with the hydraulic architecture of the leaf. The present study confirms the ‘classical’ view of the hydraulic architecture of leaves as composed by a low‐resistance component (the venation) and a high‐resistance component (the mesophyll).     

Hydraulic architecture and water flow in growing grass tillers (Festuca arundinacea Schreb.)

The water relations and hydraulic architecture of growing grass tillers (Festuca arundinacea Schreb.) are reported. Evaporative flux density, E (mmol s^?1 m^?2), of individual leaf blades was measured gravimetrically by covering or excision of entire leaf blades. Values of E were similar for mature and elongating leaf blades, averaging 2·4 mmol s^?1 m^?2. Measured axial hydraulic conductivity, K_h (mmol s^?1 mm MPa^?1), of excised leaf segments was three times lower than theoretical hydraulic conductivity (K_t) calculated using the Poiseuille equation and measurements of vessel number and diameter. K_t was corrected (K_t*) to account for the discrepancy between K_h and K_t and for immature xylem in the basal expanding region of elongating leaves. From base to tip of mature leaves the pattern of K_t* was bell‐shaped with a maximum near the sheath–blade joint (≈ 19 mmol s^?1 mm MPa^?1). In elongating leaves, immature xylem in the basal growing region led to a much lower K_t*. As the first metaxylem matured, K_t* increased by 10‐fold. The hydraulic conductances of the whole root system, (mmol s^?1 MPa^?1) and leaf blades, (mmol s^?1 MPa^?1) were measured by a vacuum induced water flow technique. and were linearly related to the leaf area downstream. Approximately 65% of the resistance to water flow within the plant resided in the leaf blade. An electric‐analogue computer model was used to calculate the leaf blade area‐specific radial hydraulic conductivity, (mmol s^?1 m^?2 MPa^?1), using , K_t* and water flux values. values decreased with leaf age, from 21·2 mmol s^?1 m^?2 MPa^?1 in rapidly elongating leaf to 7·2 mmol s^?1 m^?2 MPa^?1 in mature leaf. Comparison of and values showed that ≈ 90% of the resistance to water flow within the blades resided in the liquid extra‐vascular path. The same algorithm was then used to compute the xylem and extravascular water potential drop along the liquid water path in the plant under steady state conditions. Predicted and measured water potentials matched well. The hydraulic design of the mature leaf resulted in low and quite constant xylem water potential gradient (≈ 0·3 MPa m^?1) throughout the plant. Much of the water potential drop within mature leaves occurred within a tenth of millimetre in the blade, between the xylem vessels and the site of water evaporation within the mesophyll. In elongating leaves, the low K_t* in the basal growth zone dramatically increased the local xylem water potential gradient (≈ 2·0 MPa m^?1) there. In the leaf elongation zone the growth‐induced water potential difference was ≈ 0·2 MPa.     

Ion-mediated increase in the hydraulic conductivity of Laurel stems: role of pits and consequences for the impact of cavitation on water transport

Changes in hydraulic conductivity (K(h)) were measured in stems of Laurus nobilis L. during perfusion with KCl, NaCl or sucrose solutions. Ionic solutes induced marked increase of K(h) with respect to deionized water but sucrose had no effect. The kinetics of KCl-induced K(h) increase was measured together with changes in [K(+)] of the perfused solution. K(h) increases were paralleled by increases in the [K(+)](out)/[K(+)](in) ratio. Samples of different lengths or with increasing percentage loss of conductivity (PLC) due to xylem cavitation were tested, with the aim of increasing radial flow through intervessel pits. KCl solutions enhanced the K(h) of 12-cm-long samples with a concentration-dependent effect up to 100 mm KCl. DeltaK(h) increased from 3 to 30% in 1.5- and 12-cm-long samples, respectively and remained constant for longer samples. Increasing PLC induced an exponential increase in DeltaK(h). PLC measured with KCl solutions was significantly less than that measured with deionized water, suggesting that measurements of PLC can be affected by the composition of the perfused solution. Experiments support the hypothesis that the 'ionic effect' is mediated by physico-chemical changes of pectins of the pit membranes and raise the possibility that plants might alter the ionic composition of the xylem sap to alleviate the hydraulic impact of cavitation.     

Floral biology of the dioecious species Laurus nobilis L. (Lauraceae)

The present study examines the cytological, physiological, chemical and ecological characteristics of pollen and nectar offered by male and female flowers of the dioecious plant Laurus nobilis. The various phases of floral phenology and the insect pollinators were observed. We used cytological methods to determine anther, pollen and nectary structure. Nectar sugar composition was evaluated by HPLC. Pollen viability in time was compared with cytoplasmic and intine water content. Pollen presentation was found to be reversible by opening and closing of anther valves, determined by hydration of the mechanical layer of the anther. Pollen, covered by pollenkitt, was presented for dispersal for 3 consecutive days and during this time the intine and cytoplasm lost water and pollen viability diminished. At germination exine ruptured together with the outermost layer of the intine. Nectaries of male flowers were observed on the anther filament and on staminodes of female flowers. The nectar consisted almost entirely of sucrose and was more concentrated in male flowers. Secreted through stomata, nectar was presented in a thin layer. In the study area, the main pollinators (about half the total number of all visits) were hymenopterans. Pollen is of the recalcitrant type due to its high water content (>30%) but its viability is long-lasting because the intine is thick and stores water, keeping the cytoplasm of the vegetative cell hydrated and viable, and because anther valves may close under adverse conditions, protecting the pollen. Insects are attracted by male and female flowers similarly, males offer nectar and pollen, whilst females only nectar.     

Direct identification and quantitative determination of costunolide and dehydrocostuslactone in the fixed oil of Laurus novocanariensis by 13C-NMR spectroscopy

The fixed oil of Laurus novocanariensis (previously L. azorica) contains mostly glycerides together with minor non-saponifiable compounds. The direct identification and quantitative determination of costunolide and dehydrocostuslactone, two sesquiterpene lactones components of the oil that exhibit biological activities, is described. The analysis was carried out using 13C-NMR spectroscopy (signal acquisition with inverse gated decoupling of protons; diglyme as internal standard) without separation, derivatisation or any sample preparation.     

Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought

A comparative study on stomatal control under water deficit was conducted on grapevines of the cultivars Grenache, of Mediterranean origin, and Syrah of mesic origin, grown near Montpellier, France and Geisenheim, Germany. Syrah maintained similar maximum stomatal conductance (g_max) and maximum leaf photosynthesis (A_max) values than Grenache at lower predawn leaf water potentials, Ψ_leaf, throughout the season. The Ψ_leaf of Syrah decreased strongly during the day and was lower in stressed than in watered plants, showing anisohydric stomatal behaviour. In contrast, Grenache showed isohydric stomatal behaviour in which Ψ_leaf did not drop significantly below the minimum Ψ_leaf of watered plants. When g was plotted versus leaf specific hydraulic conductance, K_l, incorporating leaf transpiration rate and whole‐plant water potential gradients, previous differences between varieties disappeared both on a seasonal and diurnal scale. This suggested that isohydric and anisohydric behaviour could be regulated by hydraulic conductance. Pressure‐flow measurements on excised organs from plants not previously stressed revealed that Grenache had a two‐ to three‐fold larger hydraulic conductance per unit path length (K_h) and a four‐ to six‐fold larger leaf area specific conductivity (LSC) in leaf petioles than Syrah. Differences between internodes were only apparent for LSC and were much smaller. Cavitation detected as ultrasound acoustic emissions on air‐dried shoots showed higher rates for Grenache than Syrah during the early phases of the dry‐down. It is hypothesized that the differences in water‐conducting capacity of stems and especially petioles may be at the origin of the near‐isohydric and anisohydric behaviour of g.     

Leaf responses to soil water deficits: Comparative sensitivity of leaf expansion rate and leaf conductance in field-grown sunflower (Helianthus annuus L.)

The relative importance of changes in leaf expansion rate (LER) and leaf conductance (g₁) in the control of crop transpiration depends primarily on their sensitivity to soil water deficits. The aim of this paper was to quantify the responses of LER and g₁ to soil water deficits in sunflower (Helianthus annuus L.) under conditions of moderate (spring) and high (summer) evaporative demand. Soil water content, g₁, and LER were measured in dryland (DRY) and daily-irrigated (WET) crops established on a deep sandy-loam (Typic Xerofluvent) in a Mediterranean environment. There was no difference between g₁ of DRY and WET plants (p>0.20) in contrast with a highly significant difference in LER (p<0.001). Even under the harsh conditions of the summer experiment, g₁ did not respond to water deficit in a ten-day period in which LER of DRY plants was reduced to approx. 30% of that measured in WET controls. This field study indicates that g₁ plays at most a minor role in the control of sunflower transpiration in the pre-anthesis period and confirms the importance of leaf expansion in the regulation of gas exchange of expanding canopies subjected to soil water deficits.     

An anatomical assessment of branch abscission and branch-base hydraulic architecture in the endangered Wollemia nobilis

Background and Aims: The branch-base xylem structure of the endangered Wollemia nobiliswas anatomically investigated. Wollemia nobilis is probablythe only extant tree species that produces only first-orderbranches and where all branches are cleanly abscised. An investigationwas carried out to see if these unusual features might influencebranch-base xylem structure and water supply to the foliage. Methods: The xylem was sectioned at various distances along the branchbases of 6-year-old saplings. Huber values and relative theoreticalhydraulic conductivities were calculated for various regionsof the branch base. Key Results: The most proximal branch base featured a pronounced xylem constriction.The constriction had only 14–31 % (average 21 %)of the cross-sectional area and 20–42 % (average28 %) of the theoretical hydraulic conductivity of themore distal branch xylem. Wollemia nobilis had extremely lowHuber values for a conifer. Conclusions: The branch-base xylem constriction would appear to facilitatebranch abscission, while the associated Huber values show thatW. nobilis supplies a relatively large leaf area through a relativelysmall diameter ‘pipe’. It is tempting to suggestthat the pronounced decline of W. nobilis in the Tertiary isrelated to its unusual branch-base structure but physiologicalstudies of whole plant conductance are still needed.     

Hydraulic architecture of sugarcane in relation to patterns of water use during plant development*

Hydraulic conductance was measured on leaf and stem segments excised from sugarcane plants at different stages of development. Maximum transpiration rates and leaf water potential (Ψ_L) associated with maximum transpiration were also measured in intact plants as a function of plant size. Leaf specific hydraulic conductivity (L_sc) and transpiration on a unit leaf area basis (E) were maximal in plants with approximately 0.2 m² leaf area and decreased with increasing plant size. These changes in Fand L_sc were nearly parallel, which prevented φ_L in larger plants from decreasing to levels associated with substantial loss in xylem conductivity caused by embolism formation. Coordination of changes in E and leaf hydraulic properties was not mediated by declining leaf water status, since φ_L increased with plant size. Hydraulic constrictions were present at nodes and in the node-leaf sheath-leaf blade pathway. This pattern of constrictions is in accord with the idea of plant segmentation into regions differing in water transport efficiency and would tend to confine embolisms to the relatively expendable leaves at terminal positions in the pathway, thereby preserving water transport through the stem.     

Developmental patterns of above-ground hydraulic conductance in a Scots pine (Pinus sylvestris L.) age sequence

Hydraulic resistance to water flow was measured in branches and stems of Scots pine trees ranging from 7 to 59 years of age in Thetford (East Anglia, UK). On the basis of these measurements, tree above-ground conductance was calculated and related to the amount of leaf area sustained by each tree. Branches at the crown bottom had a lower proportion of sapwood area and a lower total hydraulic conductance than branches of the same diameter at the tree top. Within branches, most of the hydraulic resistance was located near the needles. Tree above-ground conductance was positively related to tree diameter and inversely related to tree height. Compared with young trees, mature trees had about 4 times less above-ground conductance per unit of leaf area. Apparently, the increase in pathway length associated with tree height growth could be only partially compensated for by the increase in conductive capacities resulting from diameter growth. We argue that this reduction may account for reported decreases of stomatal conductance with tree age. It is suggested that the increase in branchiness associated with tree maturation may represent a compensation mechanism to reduce the overall resistance to water flow in the crowns.     

麦冬、土麦冬和阔叶土麦冬叶表皮形态结构的观察

用光镜和扫描电镜观察了麦冬（Ophiopogon japonicus（L．f）Ker—Gawl.]、土麦冬（Liriope spicata Lour．）和阔叶土麦冬（L．platyphylla Wanget Tang）叶表皮显微结构、亚显微结构和角质层内表面的形态结构。结果表明,气孔主要分布于麦冬、土麦冬和阔叶土麦冬叶片的下表皮,气孔密度分别为76．4、114．3和99．8个·mm^-2;仅阔叶土麦冬叶片上表皮有少量气孔分布。3种植物的气孔器均不具有副卫细胞,并在叶脉间形成纵向气孔带。表皮细胞长方形,气孔带与非气孔带处表皮细胞的形态和大小差异较明显。麦冬气孔周围的表皮细胞平周壁具明显瘤状突起,导致气孔下陷;土麦冬气孔周围的表皮细胞平周壁呈波浪状突起,使气孔相对下陷;阔叶土麦冬气孔周围的表皮细胞平周壁基本无突起,气孔不下陷。3种植物的叶表皮均有发达的角质层和丰富的蜡质,且蜡质主要分布于下表皮气孔带处。这些结构特征可能与它们所具有的喜阳、耐阴和耐旱等特性有一定的相关性。     

Control of stomatal conductance by leaf water potential in Hymenoclea salsola (T. & G.), a desert subshrub

The role of leaf water potential in controlling stomatal conductance ( g _s) was examined in the desert subshrub Hymenoclea salsola . For plants operating at high irradiance, stomatal closure in response to high leaf-air humidity gradient ( D ) was largely reversed by soil pressurization. Stomatal re-opening eliminated, on average, 89% of the closure normally induced by high D . Transpiration rates ( E ) reached under these conditions were far higher than maximal rates normally observed at any point of the D response. In situ stem psychrometry indicated that water flux at all times conformed to a simple Ohm's-law analogy. Under conditions of high D, E increased substantially in response to soil pressurization. Stomatal regulation did not constrain E during this treatment, but did result in nearly constant minimum leaf water potentials.     

黄独(Dioscorea bulbifera L.)不同居群叶表皮微形态特征的比较观察

应用扫描电子显微镜(SEM)对15个黄独(Dioscorea bulbifera L.)居群(11个野生居群及4个栽培居群)植株的叶表皮微形态特征(包括气孔器、表皮毛和气孔周围表皮细胞的特征)进行了观察和测量,并据此编制了15个黄独居群的检索表.观察结果表明:15个黄独居群植株叶片在气孔器外拱盖内缘类型,气孔器旋转方向,气孔长轴长度及密度和深度,叶片表皮毛的有无及类型,叶肉腺毛的长、短轴长度及密度,气孔周围表皮细胞垂周壁和平周壁的形态以及平周壁表面颗粒形状等特征上均有明显差异.气孔器外拱盖内缘有光滑和浅波状2种类型;气孔器的旋转方向分为不定向旋转、左旋和右旋3种方式;气孔长轴长度和密度的变化幅度分别为18.59～31.93 μm和86～356 mm-2.叶片表皮毛均仅存在于下表皮,可分为乳突和多细胞头单细胞柄腺毛2种类型,乳突主要分布于主脉,腺毛主要分布于二级脉和叶肉;叶肉腺毛的长、短轴长度的变化幅度分别为25.00～55.00和24.86～44.29μm,大部分居群腺毛密度的变化幅度为6～29 mm-2.叶片气孔周围表皮细胞垂周壁的形状有平直、平直隆起、平直脊状隆起、弯曲、弯曲脊状隆起和弯曲拱状隆起6种类型;表皮细胞平周壁的表面纹饰有具瘤条纹和光滑条纹2种形态,其扩散方式也有2种形式:一种为不环绕气孔且四方扩散,另一种为环绕气孔2～3周后扩散但扩散方向不定;平周壁上的颗粒形状有簇晶状、屑状、密集粉状、粉状、粒状和片状6种类型.比较分析结果显示:黄独不同居群叶表皮微形态特征的多态性较为丰富,地理分布相近的居群微形态特征的相似性较高,这些特征与黄独的各种下分类群相对应.