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Boron supply into wheat (Triticum aestivum L. cv. Wilgoyne) ears whilst still enclosed within leaf sheaths.

The present study investigates whether there is significant remobilization of (10)B previously loaded in the flag and penultimate leaves into the young, actively growing ear enclosed within the sheaths of flag and penultimate leaves. It also explores whether B transport into the enclosed ear declines when air humidity in the shoot canopy increases. After 5 d (10)B labelling during the period from early to full emergence of the flag leaf, the plants were transferred into nutrient solutions containing either 10 microM (11)B or no added B for 3 d. Regardless of the subsequent B supply levels to the roots, (10)B contents in the ear continued to increase by up to 5-fold 3 d after the end of (10)B supply in the nutrient solution. During these 3 d, the ear experienced a rapid increase in biomass. However, the majority of B in the ear during the 3 d treatment period was from the newly acquired (11)B from root uptake, rather than retranslocation of (10)B previously deposited in the leaves. By comparing the relative distribution of (10)B, Rb (xylem-to-phloem transfer marker) and Sr (xylem-marker) in the ear and the flag leaf, the distribution of (10)B resembled that of Rb more than Sr. Canopy cover treatment greatly suppressed leaf transpiration and decreased the amount of newly acquired (10)B in the flag leaf and the ear, but not in the upper stem segments. The results suggest that whilst the young ear was still fully enclosed within the leaf sheaths without any significant transpiration activity, B transport into the ear is predominantly dependent on the long-distance B transport in the xylem driven by leaf transpiration and, therefore, on concurrent B uptake from the roots.

Biological Transport↗

[Anatomy of Aconitum finetianum Hand.-Mazz].

The primary xylem of plump taproot of Aconitum finetianum is the diarch with xylem in the center. In the action of the secondary structure of the herb, the meristem is first produced in the secondary phloem, then the anomalous cambium is produced in the secondary xylem forms a new and splits into anomalous structure. In the action of anomalous structure of the herb, the meristem in the secondary phloem sinks inwardly from many sites, and forms many rings with the anomalous cambium. One of these rings forms a special endodermis and the protostele turns into the schizogenous stele.

Magnoliopsida↗

The apoplastic pH of the substomatal cavity of Vicia faba leaves and its regulation responding to different stress factors.

The apoplastic pH of the substomatal cavity is an essential determinant of stomatal movement. In detached leaves of Vicia faba substomatal apoplastic pH and its dependence on external (stress) factors was investigated using a non-invasive approach: pH-microsensors were inserted into open stomata, and upon contact with the apoplastic fluid, pH was measured continuously, as apoplastic pH was challenged by changed conditions of light, atmosphere (NH(3), CO(2)), and xylem sap (abscisic acid, cyanide, fusicoccin, pH, inorganic salts). Apoplastic pH proved extremely sensitive to infiltration and local flooding, which rapidly increased the apoplastic pH by more than 1.5 pH units. Recovery from infiltration took several hours, during which light effects on the apoplastic pH were strongly impeded. This indicates that pH tests carried out under such conditions may not be representative of the undisturbed leaf. NH(3), flushed across the stomata, yielded a rapid apoplastic alkalinization from which an apoplastic buffer capacity of 2-3 mM per pH unit was calculated. Fusicoccin, fed into the xylem sap acidified the apoplast, whereas cyanide alkalized it, thus underscoring the importance of the plasma membrane H(+) pump for apoplastic pH regulation. To address the question to what extent pH was a drought signal, the effect of iso-osmotic pH changes, fed into the xylem through the petiole were tested. It is demonstrated that the apoplastic response remained below 0.1 pH per pH unit imposed, regardless of the buffer capacity. An increase in the osmolarity of the bath solution (harbouring the cut petiole) using KCl, NaCl, CaCl(2) or sorbitol alkalized the substomatal apoplast. It is suggested that pH may only act as drought signal when accompanied by elevated osmolarity.

Abscisic Acid↗

[Hydraulic characters of Acer truncatum seedlings].

The hydraulic parameters of 4 years old Acer truncatum seedlings were measured by improved flushing method under the condition of controlled drought gradients in the greenhouse. It's indicated that the changes of hydraulic parameters with stem segment functional xylem diameter could be modeled by different equations. The hydraulic conductivity was influenced by the area that stem segment located. It was higher in non-constriction area than in constriction area. The existence of constriction area was in favor of the competition between individual seedlings. Hydraulic conductivity, specific conductivity and leaf specific conductivity were proportional with functional xylem diameter and twig water potential. The leaf specific conductivity of thicker branches was far higher than that of distal twigs, which was in favor of seedlings in saving those organs with more photosynthesis devotion during drought stress. The change of Huber value of same diameter branches with twig water potential was very small before defoliation, and hence, the main source of seedling's water stress came from xylem cavitation and embolism.

Acer↗

Inhibition of lignin formation by L-alpha-aminooxy-beta-phenylpropionic acid, an inhibitor of phenylalanine ammonia-lyase.

Mungbean (Vigna radiata (L.) Wilczek) seedlings grown for 9 days on filter paper soaked with 0.3 to 1 mM L-alpha-aminooxy-beta-phenylpropionic acid (AOPP), a potent inhibitor of L-phenylalanine ammonia-lyase, had a greatly reduced anthocyanin content, and the cell walls of the xylem vessels did not stain with the phloroglucinol/HCl or safranine/astrablue reagents indicating the absence of lignin-like material. Furthermore, vanillin was detectable in nitro-benzene-oxidized lignin preparations only from control seedlings, but not from AOPP-treated seedlings. Scanning electron microscopy of hypocotyl cross sections revealed collapsed xylem vessels in seedlings grown in the presence of AOPP indicating that lignin is required for resistance against the tensile forces in the conducting cells of the xylem. AOPP enhanced the growth of cultured cells of Lonicera prolifera Rehd. while it inhibited the production of extracellular material that gave a positive reaction with phloroglucinol/HCl.

Ammonia-Lyases↗

Evidence of threshold temperatures for xylogenesis in conifers at high altitudes.

Temperature is the most important factor affecting growth at high altitudes. As trees use much of the allocated carbon gained from photosynthesis to produce branches and stems, information on the timing and dynamics of secondary wood growth is crucial to assessing temperature thresholds for xylogenesis. We have carried out histological analyses to determine cambial activity and xylem cell differentiation in conifers growing at the treeline on the eastern Alps in two sites during 2002-2004 with the aim of linking the growth process with temperature and, consequently, of defining thresholds for xylogenesis. Cambial activity occurred from May to July-August and cell differentiation from May-June to September-October. The earliest start of radial enlargement was observed in stone pine in mid-May, while Norway spruce was the last species to begin tracheid differentiation. The duration of wood formation varied from 90 to 137 days, depending on year and site, with no difference between species. Longer durations were observed in trees on the south-facing site because of the earlier onset and later ending of cell production and differentiation. The threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regressions. Xylogenesis was active when the mean daily air temperature was 5.6-8.5 degrees C and mean stem temperature was 7.2-9 degrees C. The similar thresholds among all trees suggested the existence of thermal limits in wood formation that correspond with temperatures of 6-8 degrees C that are supposed to limit growth at the treeline. Different soil temperature thresholds between sites indicated that soil temperature may not be the main factor limiting xylogenesis. This study represents the first attempt to define a threshold through comparative assessment of xylem growth and tissue temperatures in stem meristems at high altitudes.

Altitude↗

Characterization of two laccases of loblolly pine (Pinus taeda) expressed in tobacco BY-2 cells.

We previously showed that eight laccase genes (Lac 1-Lac 8) are preferentially expressed in differentiating xylem and are associated with lignification in loblolly pine (Pinus taeda) [Sato et al. (2001) J Plant Res 114:147-155]. In this study we generated transgenic tobacco suspension cell cultures that express the pine Lac 1 and Lac 2 proteins, and characterized the abilities of these proteins to oxidize monolignols. Lac 1 and Lac 2 enzymatic activities were detected only in the cell walls of transgenic tobacco cells, and could be extracted with high salt. The optimum pH for laccase activity with coniferyl alcohol as substrate was 5.0 for Lac 1 and between 5.0 and 6.0 for Lac 2. The activities of Lac 1 and Lac 2 increased as the concentration of CuSO(4) in the reaction mixtures increased in the range from 1 to 100 microM. Both enzymes were able to oxidize coniferyl alcohol and to produce dimers of coniferyl alcohol. These results are consistent with the hypothesis that Lac 1 and Lac 2 are involved in lignification in differentiating xylem of loblolly pine.

Cell Line, Transformed↗

Why are Chloris gayana leaves shorter in salt-affected plants? Analyses in the elongation zone.

Reduced hydraulic conductance calculated from growth data was suggested to be the main reason for reduced leaf expansion in salt-stressed Chloris gayana (Rhodes grass). In this work, xylem vessel cross-sections and wall enzyme activities were analysed to re-examine the effects of salinity on leaf growth in this species. Maximal segmental growth rates were 20% lower and the growth zone was 23% shorter in leaves from salinized plants than in controls; however, growth rates between 0 mm and 15 mm from the ligule were similar in both types of leaves. Xylem cross-sectional areas in this region were about 65% smaller in leaves of salinized plants, suggesting that hydraulic restrictions in the leaves of salinized plants were much higher than overall growth reductions. Extractable xyloglucan endotransglucosylase activity in this zone was twice as high in leaves of salinized plants as in leaves of controls. Nevertheless, the activity of the extracted enzyme was not affected by up to 1 M NaCl added to the reaction medium. Therefore, increased xyloglucan endotransglucosylase activity under salinity may be due to a promotion of transcription of XTH (xyloglucan endotransglucosylase/hydrolases) genes and/or translation of preformed transcripts. These results suggest that, as in drought stress, increased activity of cell wall enzymes associated with wall loosening may contribute to the maintenance of growth under saline conditions despite hydraulic restrictions.

Cell Wall↗

Root-to-shoot long-distance circulation of nicotianamine and nicotianamine-nickel chelates in the metal hyperaccumulator Thlaspi caerulescens.

Plant metal hyperaccumulator species are widely used as models to unravel the heavy metal tolerance and hyperaccumulation mechanisms. Thlaspi caerulescens is capable of tolerating and hyperaccumulating Zn, Cd, and Ni. A search for factors involved in the cellular tolerance to Ni, based on yeast screens, led to isolation of a cDNA encoding a functional nicotianamine (NA) synthase (NAS). The T. caerulescens genome appears to contain a single copy of the NAS gene named TcNAS whose expression is restricted to the leaves. The analysis of dose-response and time-course Ni treatments have revealed that the exposure to Ni triggers the accumulation of NA in the roots. Because neither TcNAS expression nor NAS activity were detected in the roots, the NA accumulation in roots is most probably the result of its translocation from the leaves. Once in the roots, NA, together with Ni, is subsequently found in the xylem, for redirection to the aerial parts. Using liquid chromatography coupled to inductively coupled plasma or electrospray ionization mass spectrometry, it has been shown that part of the Ni is translocated as a stable Ni-NA complex in the xylem sap. This circulation of NA, Ni, and NA-Ni chelates is absent in the non-tolerant non-hyperaccumulator related species T. arvense. Taken together, the results provide direct physiological and chemical evidence for NA and NA-heavy metal complex translocation in a hyperaccumulator species.

Alkyl and Aryl Transferases↗

Microarray gene expression profiling of developmental transitions in Sitka spruce (Picea sitchensis) apical shoots.

The apical shoot drives the yearly new stem growth of conifer trees, is the primary site for the establishment of chemical and physical defences, and is important in establishing subsequent perennial growth. This organ presents an interesting developmental system, with growth and development progressing from a meristematic tip through development of a primary vascular system, to a base with fully differentiated and lignified secondary xylem on the inside and bark tissue with constitutive defence structures such as resin, polyphenolic phloem parenchyma cells, and sclereids on the outside. A spruce (Picea spp.) microarray containing approximately 16.7K unique cDNAs was used to study transcript profiles that characterize the developmental transition in apical shoots of Sitka spruce (Picea sitchensis) from their vegetative tips to their woody bases. Along with genes involved in cell-wall modification and lignin biosynthesis, a number of differentially regulated genes encoding protein kinases and transcription factors with base-preferred expression patterns were identified, which could play roles in the formation of woody tissues inside the apical shoot, as well as in regulating other developmental transitions associated with organ maturation. Preferential expression of known conifer defence genes, genes encoding defence-related proteins, and genes encoding regulatory proteins was observed at the apical shoot tip and in the green bark tissues at the apical shoot base, suggesting a commitment to constitutive defence in the apical shoot that is co-ordinated with rapid development of secondary xylem.

Antigens, Plant↗

Hydraulic architecture correlates with bud organogenesis and primary shoot growth in beech (Fagus sylvatica).

In beech (Fagus sylvatica L.), the number of leaf primordia preformed in the buds determines the length and the type (long versus short) of annual growth units, and thus, branch growth and architecture. We analyzed the correlation between the number of leaf primordia and the hydraulic conductance of the vascular system connected to the buds. Terminal buds of short growth units and axillary buds of long growth units on lower branches of mature trees were examined. Buds with less than four and more than five leaf primordia formed short and long growth units, respectively. Irrespective of the type of growth unit the bud was formed on, the occurrence of a large number of leaf primordia was associated with high xylem hydraulic conductance. Xylem conductance was correlated to the area of the outermost annual ring. These results suggest that organogenesis and primary growth in buds correlates with secondary growth of the growth units and thus with their hydraulic architecture. Possible causal relationships between the variables are discussed.

Carbon Isotopes↗

Hydraulic conductivity, photosynthesis and leaf water balance in six evergreen woody species from fall to winter.

To confirm that freeze-thaw embolism is a primary stress for evergreen woody species in winter, hydraulic conductivity, photosynthesis and leaf water potential were measured during fall and winter in trees growing in a cool temperate zone (Nikko) and in a warm temperate zone (Tokyo). We examined two evergreen conifers that naturally occur in the cool temperate zone (Abies firma Siebold & Zucc. and Abies homolepis Siebold & Zucc.), and four evergreen broad-leaved woody species that are restricted to the warm temperate zone (Camellia japonica L., Cinnamomum camphora (L.) J. Presl, Ilex crenata Thunb. and Quercus myrsinaefolia Blume). In Tokyo, where no freeze-thaw cycles of xylem sap occurred, hydraulic conductivity, photosynthesis and water balance remained constant during the experimental period. In Nikko, where there were 38 daily freeze-thaw cycles by February, neither of the tracheid-bearing evergreen conifers showed xylem embolism or leaf water deficits. Similarly, the broad-leaved evergreen trees with small-diameter vessels did not exhibit severe embolism or water deficits and maintained CO(2) assimilation even in January. In contrast, the two broad-leaved evergreen trees with large-diameter vessels showed significantly reduced hydraulic conductivity and shoot die-back in winter. We conclude that freeze-thaw embolism restricts evergreen woody species with large-diameter vessels to the warm temperate zone, whereas other stresses limit the distribution of broad-leaved trees, that have small-diameter vessels, but which are restricted to the warm temperate zone.

Abies↗

Influences of environmental factors on the radial profile of sap flux density in Fagus crenata growing at different elevations in the Naeba Mountains, Japan.

Sap flux density was measured continuously during the 1999 and 2000 growing seasons by the heat dissipation method in natural Fagus crenata Blume (Japanese beech) forests growing between 550 and 1600 m on the northern slope of the Kagura Peak of the Naeba Mountains, Japan. Sap flux density decreased radially toward the inner xylem and the decrease was best expressed in relation to the number of annual rings from the cambium, or in relation to the relative depth between the cambium and the trunk center, rather than as a function of absolute depth. The relative influences of radiation, vapor pressure deficit and soil water on sap flux density during the growing season were similar for the outer and inner xylem, and at all sites. Measurements of soil water content and water potential at a depth of 0.25 m demonstrated that sap flux density responded similarly and sensitively to water potential changes in this soil layer, despite large differences in rooting depth at different elevations, localizing one important control point in the functioning of this forest ecosystem. Identification of the relative influences of radiation, vapor pressure deficit and drying of the upper soil layer on sap flux density provides a framework for in-depth analysis of the control of transpiration in Japanese beech forests. In addition, the finding that the same general controls are operating on sap flux density despite climate gradients and large differences in overall forest stand structure will enhance understanding of water use by forests along elevation gradients.

Altitude↗

Transpiration-induced axial and radial tension gradients in trunks of Douglas-fir trees.

We determined the axial and radial xylem tension gradients in trunks of young Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Axial specific conductivity (k(s-a)) and sap flux density (Js) were measured at four consecutive depths within the sapwood at a stem height of 1 m. By definition, at a given position in the bole, Js is a function not only of k(s-a) but also of the driving force for water movement. The Js:k(s-a) ratio was therefore used to estimate axial tension gradients and the radial gradients at a stem height of 1 m were calculated from the differences in axial tension gradients at each depth. Tracheid lumen diameter and tracheid length were used to predict differences in k(s-a) and its divergence from the theoretical k(s-a) determined by the Hagen Poisseuille equation. The ratio of k(s-a) (determined in the laboratory) to Js (measured in the field) varied with depth in the sapwood, resulting in non-uniform axial and radial tension gradients from inner to outer sapwood. Transpiration-induced axial tension gradients were in the range of 0.006-0.01 MPa m(-1) excluding the gravitational tension gradient. At a stem height of 1 m, radial tension gradients were in the range of 0.15-0.25 MPa m(-1) and were lower in the middle sapwood than in the inner or outer sapwood. Axial tension gradients were 44-50% higher in the outer sapwood than in the inner sapwood. At a stem height of 1 m, radial Js, calculated on the basis of radial tension gradients and measured radial specific conductivity (k(s-r)), was about two orders of magnitude smaller than axial Js. Our findings indicate that large radial tension gradients occur in the sapwood and clarify the role played by xylem k(s-a) and k(s-r) in determining in situ partitioning of Js in the axial and radial directions.

Circadian Rhythm↗

Interactions between MUR10/CesA7-dependent secondary cellulose biosynthesis and primary cell wall structure.

Primary cell walls are deposited and remodeled during cell division and expansion. Secondary cell walls are deposited in specialized cells after the expansion phase. It is presently unknown whether and how these processes are interrelated. The Arabidopsis (Arabidopsis thaliana) MUR10 gene is required for normal primary cell wall carbohydrate composition in mature leaves as well as for normal plant growth, hypocotyl strength, and fertility. The overall sugar composition of young mur10 seedlings is not significantly altered; however, the relative proportion of pectin side chains is shifted toward an increase in 1 --> 5-alpha-arabinan relative to 1 --> 4-beta-galactan. mur10 seedlings display reduced fucogalactosylation of tightly cell wall-bound xyloglucan. Expression levels of genes encoding either nucleotide sugar interconversion enzymes or glycosyl transferases, known to be involved in primary and secondary cell wall biosynthesis, are generally unaffected; however, the CesA7 transcript is specifically suppressed in the mur10-1 allele. The MUR10 locus is identical with the CesA7 gene, which encodes a cellulose catalytic subunit previously thought to be specifically involved in secondary cell wall formation. The xylem vessels in young mur10 hypocotyls are collapsed and their birefringence is lost. Moreover, a fucogalactosylated xyloglucan epitope is reduced and a 1 --> 5-alpha-arabinan epitope increased in every cell type in mur10 hypocotyls, including cells that do not deposit secondary walls. mur10 also displays altered distribution of an arabinogalactan-protein epitope previously associated with xylem differentiation and secondary wall thickening. This work indicates the existence of a mechanism that senses secondary cell wall integrity and controls biosynthesis or structural remodeling of primary cell walls and cellular differentiation.

Alleles↗

Evidence that differential gene expression between the halophyte, Thellungiella halophila, and Arabidopsis thaliana is responsible for higher levels of the compatible osmolyte proline and tight control of Na+ uptake in T. halophila.

Salt-sensitive glycophytes and salt-tolerant halophytes employ common mechanisms to cope with salinity, and it is hypothesized that differences in salt tolerance arise because of changes in the regulation of a basic set of salt tolerance genes. We explored the expression of genes involved in two key salt tolerance mechanisms in Arabidopsis thaliana and the halophytic A. thaliana relative model system (ARMS), Thellungiella halophila. Salt overly sensitive 1 (SOS1) is a plasma membrane Na+/H+ antiporter that retrieves and loads Na+ ions from and into the xylem. Shoot SOS1 transcript was more strongly induced by salt in T. halophila while root SOS1 was constitutively higher in unstressed T. halophila. This is consistent with a lower salt-induced rise in T. halophila xylem sap Na+ concentration than in A. thaliana. Thellungiella halophila contained higher unstressed levels of the compatible osmolyte proline than A. thaliana, while under salt stress, T. halophila accumulated more proline mainly in shoots. Expression of the A. thaliana ortholog of proline dehydrogenase (PDH), involved in proline catabolism, was undetectable in T. halophila shoots. The PDH enzyme activity was lower and T. halophila seedlings were hypersensitive to exogenous proline, indicating repression of proline catabolism in T. halophila. Our results suggest that differential gene expression between glycophytes and halophytes contributes to the salt tolerance of halophytes.

Anthocyanins↗

Nutrient availability constrains the hydraulic architecture and water relations of savannah trees.

Leaf and whole plant-level functional traits were studied in five dominant woody savannah species from Central Brazil (Cerrado) to determine whether reduction of nutrient limitations in oligotrophic Cerrado soils affects carbon allocation, water relations and hydraulic architecture. Four treatments were used: control, N additions, P additions and N plus P additions. Fertilizers were applied twice yearly, from October 1998 to March 2004. Sixty-three months after the first nutrient addition, the total leaf area increment was significantly greater across all species in the N- and the N + P-fertilized plots than in the control and in the P-fertilized plots. Nitrogen fertilization significantly altered several components of hydraulic architecture: specific conductivity of terminal stems increased with N additions, whereas leaf-specific conductivity and wood density decreased in most cases. Average daily sap flow per individual was consistently higher with N and N + P additions compared to the control, but its relative increase was not as great as that of leaf area. Long-term additions of N and N + P caused midday PsiL to decline significantly by a mean of 0.6 MPa across all species because N-induced relative reductions in soil-to-leaf hydraulic conductance were greater than those of stomatal conductance and transpiration on a leaf area basis. Phosphorus-fertilized trees did not exhibit significant changes in midday PsiL. Analysis of xylem vulnerability curves indicated that N-fertilized trees were significantly less vulnerable to embolism than trees in control and P-fertilized plots. Thus, N-induced decreases in midday PsiL appeared to be almost entirely compensated by increases in resistance to embolism. Leaf tissue water relations characteristics also changed as a result of N-induced declines in minimum PsiL: osmotic potential at full turgor decreased and symplastic solute content on a dry matter basis increased linearly with declining midday PsiL across species and treatments. Despite being adapted to chronic nutrient limitations, Cerrado woody species apparently have the capacity to exploit increases in nutrient availability by allocating resources to maximize carbon gain and enhance growth. The cost of increased allocation to leaf area relative to water transport capacity involved increased total water loss per plant and a decrease in minimum leaf water potentials. However, the risk of increased embolism and turgor loss was relatively low as xylem vulnerability to embolism and leaf osmotic characteristics changed in parallel with changes in plant water status induced by N fertilization.

Climate↗

Anisohydric but isohydrodynamic: seasonally constant plant water potential gradient explained by a stomatal control mechanism incorporating variable plant hydraulic conductance.

Isohydric and anisohydric regulations of plant water status have been observed over several decades of field, glasshouse and laboratory studies, yet the functional significance and mechanism of both remain obscure. We studied the seasonal trends in plant water status and hydraulic properties in a natural stand of Eucalyptus gomphocephala through cycles of varying environmental moisture (rainfall, groundwater depth, evaporative demand) in order to test for isohydry and to provide physiological information for the mechanistic interpretation of seasonal trends in plant water status. Over a 16 month period of monitoring, spanning two summers, midday leaf water potential (psi(leaf)) correlated with predawn psi(leaf), which was correlated with water table depth below ground level, which in turn was correlated with total monthly rainfall. Eucalyptus gomphocephala was therefore not seasonally isohydric. Despite strong stomatal down-regulation of transpiration rate in response to increasing evaporative demand, this was insufficient to prevent midday psi(leaf) from falling to levels below -2 MPa in the driest month, well into the region likely to induce xylem air embolisms, based on xylem vulnerability curves obtained in the study. However, even though midday psi(leaf) varied by over 1.2 MPa across seasons, the hydrodynamic (transpiration-induced) water potential gradient from roots to shoots (delta psi(plant)), measured as the difference between predawn and midday psi(leaf), was relatively constant across seasons, averaging 0.67 MPa. This unusual pattern of hydraulic regulation, referred to here as isohydrodynamic, is explained by a hydromechanical stomatal control model where plant hydraulic conductance is dependent on transpiration rate.

Plants↗