Total Daily Water Use Research Articles

ӘРТҮРЛІ ТӘСІЛДЕРМЕН СУҒАРУ КЕЗІНДЕГІ БҰРЫШ ЖӘНЕ БАКЛАЖАН ДАҚЫЛДАРЫНЫҢ СУДЫ ПАЙДАЛАНУ КОЭФФИЦИЕНТІН ЗЕРТТЕУ

The scientific article presents the results of a study of the coefficient of water use of pepper and eggplant crops when watering in various ways. Water agriculture consumes a large amount of water. In order to save water supplied to agricultural crops, effective irrigation methods are used. The water use of crops varies and depends on various indicators. Pepper and eggplant cultures obtained as a result of research work are among the most commonly grown crops. In recent years, these crops have been drip-watered in an open field and greenhouse. In the research work, pepper and eggplant cultures were grown through seedlings in an open field, they were systematized and watered drip. Before determining the water use coefficient of vegetable crops, their total water use and average daily water use are determined. Total water use and average daily water use of pepper and eggplant crops were low in the drip irrigation variant. It was found that the height of vegetable crops in the drip irrigation variant was higher than +4+5 cm. In the variant of systematic irrigation, the coefficient of water use of vegetable crops was 40 m3/c in the corner, in the variant of drip irrigation-21 m3/c, in the variant of drip irrigation, this indicator was below 20.8 m3/c in the corner, in the variant of drip irrigation-11.2 m3/с. In the drip irrigation variant, the water use coefficient of crops was lower than 9.8-19 m3/c (47-52%).

Analysis of institutional daily domestic water consumption dynamics due to COVID-19 pandemic, a case study of Adama Science and Technology University

The coronavirus pandemic outbreak is constantly changing the way of people’s life. To minimize or stop the transmission of the virus, several different measures have been taken by countries worldwide and including in Ethiopia. The World Health Organization (WHO), the Ethiopian Ministry of Health, and the Ethiopian Public Health Institute recommended frequent hand wash with soap and water as one of the prevention measures for the transmission of the COVID-19 pandemic. Conversely, the provision of water supply, sanitation, and hygiene is the main problem in developing countries. After the COVID-19 outbreak, the water related challenges exacerbated the situation before the pandemic existed. Accordingly, the additional supply of water is required to sufficiently provide water for hand wash. This paper mainly addresses how much additional water is required daily for domestic consumption after the coronavirus pandemic by educational institutions, Adama Science and Technology University as a case study. The data was collected with a questionnaire by the randomly selected respondents and analyzed by SPSS (Statistical package for social sciences).The analysis shown that the consumption of water has been changed after the Corona virus (COVID-19) pandemic existed. The highest percentage of the responses to the questionnaire, more than 35% require water (2–5) liters per day and more than 60% of the respondents need additional water up to 5 L per day after the pandemic existed. Based on the analysis, total daily water use by the institute before the COVID-19 existed was 24.2 m3/day. These amounts of water not sufficiently satisfy the daily water requirements of the institution even before the pandemic. After the pandemic existed, the water consumption drastically increased to 29 m3/day. Furthermore, the study recommends the provision of additional water supply sources or water demand management to reduce the impact of the pandemic on water availability.

Evaluation of seasonal patterns of hydraulic redistribution in a humid subtropical area, East China

AbstractAntecedent soil moisture or soil moisture status has a great impact on hydrological processes. Hydraulic redistribution (HR), a widely observed phenomenon, is defined as water distributed (typically at night) from moist soil to drier soil via plant roots, which plays an important role in soil moisture replenishment. Knowledge on seasonal patterns of HR and on the relationship between HR and soil water use is not fully understood. We investigated temporal variations in HR and total daily water use (Δθ) at stands of camphor and peach by monitoring soil moisture content in a humid region in eastern China. HR at the three locations reached its maximum values in summer (0.68 mm d−1 to 1.15 mm d−1) at depths of 15 cm and 35 cm. Redistributed water replenished 41% of water depleted in the soil at a 5–45‐cm depth. Interestingly, normalized HR (i.e., HR/Δθ) showed a constant pattern during the growing season implying it is independent of seasonal climate alterations. This also indicated that HR had a stable effect on the replenishment of daily water use. Positive linear relationships between HR and Δθ were found at three measuring locations (camphor: R2 = .35, p < .01; peach1: R2 = .57, p < .01; peach2: R2 = .63, p < .01), suggesting a relatively stable inherent linkage between HR and Δθ. This study suggested that hydrological processes involving soil moisture status or antecedent soil moisture, needs to take the HR effect into account across timescales from intraday to seasonal.

Hydraulic redistribution and its contribution to water retention during short-term drought in the summer rainy season in a humid area

Hydraulic redistribution (HR) in sites with shallow soils and its response to random droughts during the summer rainy season in humid areas remains poorly understood. We investigated the soil moisture dynamics, HR and water buffer of shallow soils by monitoring the soil moisture content at multiple depths during a random drought event in the summer rainy season, in typical forestland and orchard in a subtropical area. Soil moisture sensors were installed at depths of 5, 10, 15, 20 and 30 cm at three sites (forest, peach1, and peach2). The total and net daily water use and water buffer capacity of the root zone were defined and calculated. During the short-term drought, the HR of soil moisture occurred at all sites, suggesting that it could be an important process in both forest and orchard land in response to short-term drought. HR occurred at both daily and multiple-day timescales in the shallow soils. The net daily water use had less difference between the sites than the total daily water use. Principal component analysis of the total and net daily water use showed only evident clustering characteristics for the total daily water use, indicating an important plant-soil interaction effect. The depth-averaged magnitude of daily HR varied from 0.385 mm (0.0077 m3 m−3) at the forest site to 0.725 mm (0.0145 m3 m−3) at the peach1 site. The redistributed water replenished over 75% of the water depleted from the shallow soil. In particular, soil moisture was better retained in 15–20 cm root zone than in other soil layers. Furthermore, the water buffer capacity of the forest site was not higher than those of the two peach sites (where land use converted from forest), indicating that land use conversion does not necessarily weaken soil water retention. The study results highlighted that HR has a significant influence on water recharge and water retention in humid area, thus benefiting plant drought tolerance and total water utilization. This study provides more insights to evaluate HR’s effect on water retention and enhance our understanding of HR in shallow soils during random droughts in the summer rainy season in humid areas.

Consequences of nocturnal water loss: a synthesis of regulating factors and implications for capacitance, embolism and use in models.

Total daily water use is a key factor influencing the growth of many terrestrial plants, and reflects both day-time and nocturnal water fluxes. However, while nocturnal sap flow (En) and stomatal conductance (gs,n) have been reported across a range of species, ecosystems and microclimatic conditions, the regulation of these fluxes remains poorly understood. Here, we present a framework describing the role of abiotic and biotic factors in regulating En and gs,n highlighting recent developments in this field. Across ecosystems, En and gs,n generally increased with increasing soil water content and vapor pressure deficit, but the interactive effects of these factors and the potential roles of wind speed and other abiotic factors remain unclear. On average, gs,n and En are higher in broad-leaved compared with needle-leaved plants, in C3 compared with C4 plants, and in tropical compared with temperate species. We discuss the impacts of leaf age, elevated [CO2] and refilling of capacitance on night-time water loss, and how nocturnal gs,n may be included in vegetation models. Younger leaves may have higher gs,n than older leaves. Embolism refilling and recharge of capacitance may affect sap flow such that total plant water loss at night may be less than estimated solely from En measurements. Our estimates of gs,n for typical plant functional types, based on the published literature, suggest that nocturnal water loss may be a significant fraction (10-25%) of total daily water loss. Counter-intuitively, elevated [CO2] may increase nocturnal water loss. Assumptions in process-based ecophysiological models and dynamic global vegetation models that gs is zero when solar radiation is zero are likely to be incorrect. Consequently, failure to adequately consider nocturnal water loss may lead to substantial under-estimation of total plant water use and inaccurate estimation of ecosystem level water balance.

Plant water status and hydraulic conductance during flowering in the southern California coastal sage shrub Salvia mellifera (Lamiaceae)

Plant water status during flowering is important for plant reproduction, but the physiology of floral water use is not well understood. We investigated plant water status in relation to leaf and floral physiology in naturally occurring individuals of a semiarid shrub, Salvia mellifera E. Greene. We measured stomatal (g(s)) and corolla (g(c)) conductance to water vapor, transpiration from leaves (E(leaf)) and corollas (E(corolla)), leaf-specific hydraulic conductance (K(H)), bulk shoot water potential (Ψ(shoot)), and shoot water content on irrigated and control plants to analyze whether water was limiting to leaf and floral water use. Experimental irrigation caused a 203% increase in soil moisture content, a 20% increase in predawn Ψ(shoot), a 29% increase in midday Ψ(shoot), and a 92% increase in K(H). Floral and leaf gas exchange did not respond significantly to water addition, indicating that rates were at seasonal maxima and not limited by water availability. Total daily water use by corollas was ∼20% of total shoot water use. There were no significant differences in total daily shoot water use with water addition. Mean shoot water content (5.07 g) was close to mean daily shoot water use (6.71 g), indicating that the equivalent of total shoot water content turned over every 0.76 d. These results demonstrate that although irrigation improved whole-plant hydraulic conductance, gas exchange was not limited by water availability. Additionally, the high water use of flowers in this species might limit future flowering and reproductive success during dry years.

Evaluation of sap flow methods to determine water use by cultivated palms

Sustainable management of water resources allocated to palm crops requires reliable tools for measuring palm water use. Thermometric sap flow methods developed for woody dicot plants hold great potential for use in palms, but there have been few investigations to determine whether such methods can be used successfully in ‘woody’ monocots. Here, we evaluate two sap flow methods for measuring whole-plant water use by potted cocos palms ( Syagrus romanzoffiana): the Heat Ratio Method (HRM) and the Compensation Heat Pulse Method (CHPM). Measurements of whole-plant water use from the HRM and the CHPM were compared to gravimetric measurements acquired from an electronic balance. Of the two methods, the HRM gave the most accurate results and it most precisely described patterns of transpiration with respect to environmental conditions and leaf morphology. Estimates of total daily water use from the HRM and the balance were highly correlated ( R 2 = 0.92; P < 0.0001) and very near to a 1:1 relationship—an excellent result given the potential for error associated with each method. As expected, the CHPM was seriously limited at low flow rates, but it agreed well with the HRM at higher flow rates (heat pulse velocity >4 cm h −1). Anatomical investigations revealed that vascular bundles in measured palm fronds were evenly distributed and the distance between bundles was comparable in scale to the diameter of probes of HRM sensors, and most likely an order of magnitude smaller than the probable zone of thermal influence for HRM measurements. This contention was supported by results of mathematical modeling suggesting that HRM heat pulse velocities in palm-like sapwood remain largely unaffected by increases in heterogeneity caused by larger xylem vessels and wider interstitial tissues. Although wounding models for heat pulse sap flow methods have not historically catered for the effects of discrete vascular bundles, they are sufficiently approximate given our empirical validations, and we conclude that palm sapwood is thermally homogenous enough for the HRM and the CHPM to be used without modification. Overall, we present strong evidence that heat pulse methods such as the HRM and the CHPM can be used successfully in woody monocots. Finally, we note that the CHPM is likely to perform better in cases of very high flow rates, while the HRM provides more comprehensive monitoring of the usual range of flow rates including at night, early morning and during water deficit.

The effect of in-stream activities on the Njoro River, Kenya. Part I: Stream flow and chemical water quality

For shallow streams in sub-Saharan Africa, in-stream activities could be described as the actions by people and livestock, which take place within or besides stream channels. This study examined the nature of in-stream activities along a rural stream in Kenya and established the inequality in water allocation for various livelihood needs, as well as the negative impact they have on dry weather stream flow and chemical water quality. Seven locations along the stream were studied in wet and dry weather of 2006. Enumeration consisted of making head counts of people and livestock and tallying visitors at hourly intervals from 6 a.m. to 7 p.m. To estimate water abstraction, filled containers of known volume were counted and the stream was sampled to examine the impact on water quality. Water samples were obtained upstream and downstream of in-stream activities before (6 a.m.) and during (11 a.m., 6 p.m.) activities. Samples were analyzed for suspended solids, turbidity, BOD 5, total nitrogen and total phosphorus. The daily total abstraction at the middle reaches during dry weather was 120–150 m 3 day −1. More than 60% of abstraction was done by water vendors. Vended water from the stream was sold at US$ 3.5–7.5 per m 3 and vendors earned between US$ 3–6 a day. Abstracted water contributed approximately 40–60% of the total daily consumptive water use in the riparian area during dry weather but >30% of the morning stream flow was abstracted thereby upsetting stream flow in the lower reaches. The daily total water abstraction correlated positively ( R 2, 0.98) and significantly ( p < 0.05) with the daily total human visit, which was diurnally periodic with two peaks, occurring between 9 a.m. and 10 a.m. and from 4 p.m. to 5 p.m. This diurnal pattern of visits and the corresponding in-stream activities affected water quality. In particular, suspended solids, turbidity and BOD 5 levels increased significantly ( p < 0.05) downstream during in-stream activities. It was concluded that the positive contribution of in-stream activities, in particular, water abstraction to livelihoods and the daily water needs was overshadowed by the apparent disregard of the impact on stream flow and water quality. Therefore, measures are required to control in-stream activities along the stream but authorities should be mindful of the implications of any management strategy on the livelihoods of the riparian inhabitants.

A comparison of daily water use estimates derived from constant-heat sap-flow probe values and gravimetric measurements in pot-grown saplings

Use of Granier-style heat dissipation sensors to measure sap flow is common in plant physiology, ecology and hydrology. There has been concern that any change to the original Granier design invalidates the empirical relationship between sap flux density and the temperature difference between the probes. Here, we compared daily water use estimates from gravimetric measurements with values from variable length heat dissipation sensors, which are a relatively new design. Values recorded during a one-week period were compared for three large pot-grown saplings of each of the tropical trees Pseudobombax septenatum (Jacq.) Dugand and Calophyllum longifolium Willd. For five of the six individuals, P values from paired t-tests comparing the two methods ranged from 0.12 to 0.43 and differences in estimates of total daily water use over the week of the experiment averaged < 3%. In one P. septenatum sapling, the sap flow sensors underestimated water use relative to the gravimetric measurements. This discrepancy could have been associated with naturally occurring gradients in temperature that reduced the difference in temperature between the probes, which would have caused the sensor method to underestimate water use. Our results indicate that substitution of variable length heat dissipation probes for probes of the original Granier design did not invalidate the empirical relationship determined by Granier between sap flux density and the temperature difference between probes.

Nighttime transpiration in woody plants from contrasting ecosystems

It is commonly assumed that transpiration does not occur at night because leaf stomata are closed in the dark. We tested this assumption across a diversity of ecosystems and woody plant species by various methods to explore the circumstances when this assumption is false. Our primary goals were: (1) to evaluate the nature and magnitude of nighttime transpiration, E(n), or stomatal conductance, g(n); and (2) to seek potential generalizations about where and when it occurs. Sap-flow, porometry and stable isotope tracer measurements were made on 18 tree and eight shrub species from seven ecosystem types. Coupled with environmental data, our findings revealed that most of these species transpired at night. For some species and circumstances, nighttime leaf water loss constituted a significant fraction of total daily water use. Our evidence shows that E(n) or g(n) can occur in all but one shrub species across the systems we investigated. However, under conditions of high nighttime evaporative demand or low soil water availability, stomata were closed and E(n) or g(n) approached zero in eleven tree and seven shrub species. When soil water was available, E(n) or g(n) was measurable in these same species demonstrating plasticity for E(n) or g(n). We detected E(n) or g(n) in both trees and shrubs, and values were highest in plants from sites with higher soil water contents and in plants from ecosystems that were less prone to atmospheric or soil water deficits. Irrespective of plant or ecosystem type, many species showed E(n) or g(n) when soil water deficits were slight or non-existent, or immediately after rainfall events that followed a period of soil water deficit. The strongest relationship was between E(n) or g(n) and warm, low humidity and (or) windy (> 0.8 m s(-1)) nights when the vapor pressure deficit remained high (> 0.2 kPa in wet sites, > 0.7 kPa in dry sites). Why E(n) or g(n) occurs likely varies with species and ecosystem type; however, our data support four plausible explanations: (1) it may facilitate carbon fixation earlier in the day because stomata are already open; (2) it may enhance nutrient supply to distal parts of the crown when these nutrients are most available (in wet soils) and transport is rapid; (3) it may allow for the delivery of dissolved O(2) via the parenchyma to woody tissue sinks; or (4) it may occur simply because of leaky cuticles in older leaves or when stomata cannot close fully because of obstructions from stomatal (waxy) plugs, leaf endophytes or asymmetrical guard cells (all non-adaptive reasons). We discuss the methodological, ecophysiological, and theoretical implications of the occurrence of E(n) or g(n) for investigations at a variety of scales.

Significant transpirational water loss occurs throughout the night in field-grown tomato.

Incomplete stomatal closure at night can result in substantial water loss at times when photosynthetic carbon gain is not occurring in C3 and C4 plant species. To investigate the magnitude of nighttime water loss for a crop species in the field, measurements of nighttime water loss by tomato (Lycopersicon esculentum Mill. cv. Heinz 8892) were made by three methods: a field-scale lysimeter and two leaf-level instruments, an automated viscous flow porometer and a portable photosynthesis system. The portable photosynthesis system indicated nighttime transpiration of 10% of maximal daytime transpiration and the viscous flow porometer demonstrated partially open stomata. Integrated crop water loss during the dark, non-photosynthetic hours measured on the lysimeter was 3-10.8% of total daily water loss. In the glasshouse, a survey of closely related wild and cultivated tomato species showed that under ambient conditions nighttime transpiration varied within and among species and was 8-33% of maximal daytime transpiration. Implications of such a substantial fraction of total daily crop water use occurring during the night are significant in agronomic, environmental, and economic terms. Further, variation within and among species in nighttime water loss has implications for breeding to improve crop water use efficiency.

Dynamics of transpiration, sap flow and use of stored water in tropical forest canopy trees.

In large trees, the daily onset of transpiration causes water to be withdrawn from internal storage compartments, resulting in lags between changes in transpiration and sap flow at the base of the tree. We measured time courses of sap flow, hydraulic resistance, plant water potential and stomatal resistance in co-occurring tropical forest canopy trees with trunk diameters ranging from 0.34-0.98 m, to determine how total daily water use and daily reliance on stored water scaled with size. We also examined the effects of scale and tree hydraulic properties on apparent time constants for changes in transpiration and water flow in response to fluctuating environmental variables. Time constants for water movement were estimated from whole-tree hydraulic resistance (R) and capacitance (C) using an electric circuit analogy, and from rates of change in water movement through intact trees. Total daily water use and reliance on stored water were strongly correlated with trunk diameter, independent of species. Although total daily withdrawal of water from internal storage increased with tree size, its relative contribution to the daily water budget (approximately 10%) remained constant. Net withdrawal of water from storage ceased when upper branch water potential corresponded to the sapwood water potential (Psi(sw)) at which further withdrawal of water from sapwood would have caused Psi(sw) to decline precipitously. Stomatal coordination of vapor and liquid phase resistances played a key role in limiting stored water use to a nearly constant fraction of total daily water use. Time constants for changes in transpiration, estimated as the product of whole- tree R and C, were similar among individuals (~0.53 h), indicating that R and C co-varied with tree size in an inverse manner. Similarly, time constants estimated from rates of change in crown and basal sap flux were nearly identical among individuals and therefore independent of tree size and species.

Reliance on stored water increases with tree size in three species in the Pacific Northwest.

In tall old forests, limitations to water transport may limit maximum tree height and reduce photosynthesis and carbon sequestration. We evaluated the degree to which tall trees could potentially compensate for hydraulic limitations to water transport by increased use of water stored in xylem. Using sap flux measurements in three tree species of the Pacific Northwest, we showed that reliance on stored water increases with tree size and estimated that use of stored water increases photosynthesis. For Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), water stored in xylem accounted for 20 to 25% of total daily water use in 60-m trees, whereas stored water comprised 7% of daily water use in 15-m trees. For Oregon white oak (Quercus garryana Dougl. ex Hook.), water stored in xylem accounted for 10 to 23% of total daily water use in 25-m trees, whereas stored water comprised 9 to 13% of daily water use in 10-m trees. For ponderosa pine (Pinus ponderosa Dougl. ex Laws.), water stored in xylem accounted for 4 to 20% of total daily water use in 36-m trees, whereas stored water comprised 2 to 4% of daily water use in 12-m trees. In 60-m Douglas-fir trees, we estimated that use of stored water supported 18% more photosynthesis on a daily basis than would occur if no stored water were used, whereas 15-m Douglas-fir trees gained 10% greater daily photosynthesis from use of stored water. We conclude that water storage plays a significant role in the water and carbon economy of tall trees and old forests.

Canopy Position and Leaf Age Affect Stomatal Response and Water Use of Citrus

Summary We investigated the responses of citrus stomata to leaf age, leaf position, canopy size and local microclimate. Large differences existed in the stomatal response due to leaf age within a 15-year-old canopy. Much smaller differences due to leaf age were evident in 7-year-old and 2-year-old trees. Likewise, position in the canopy played a significant role in determining stomatal response within a 15-year-old tree, but had little influence in the smaller trees. Stomatal conductance data were in good agreement with values calculated using a previously published stomatal-response function based on incident photosynthetically active radiation and the vapor pressure deficit of air. The model of stomatal conductance was used to calculate total daily plant water use of 2-year-old trees based on the Penman-Monteith model. Calculated values of plant water use were close to actual values for these 2-year-old trees as measured using weighing lysimeters. These results confirm the robustness of the Penman-Monte...

Influence of rate of change in stomatal conductance to fluctuating irradiance on estimates of daily water use by Pinus taeda leaves.

Pinus taeda L. stomata respond slowly to changes in irradiance. Because incident irradiance on a leaf varies constantly, the rate of change in stomatal conductance to fluctuating irradiance may have a large effect on plant water use. We estimated total daily water use of Pinus taeda foliage for 10 days with very different irradiance patterns, assuming that rates of stomatal opening and closing were similar. To determine how the most extreme imbalance in rates of stomatal opening and closing affects estimates of water use, we also estimated total daily water use assuming instantaneous stomatal opening and a realistic rate of stomatal closing. Total daily water use was calculated by summing estimates of transpiration based on irradiance and vapor pressure deficit measured every minute at locations atop and within a canopy. Estimates of total daily water use calculated on the basis of realistic rates of stomatal opening and closing were similar to estimates calculated assuming instantaneous stomatal change (mean difference between methods of calculation was less than 0.2%). Estimates of total daily water use assuming instantaneous stomatal opening and a realistic rate of closing differed from estimates of total daily water use based on similar rates of stomatal opening and closing. The discrepancy was greater within the canopy (mean difference 6%) than at the top of the canopy (mean difference 1%). Calculation of mean daily conductance from mean daily irradiance, without accounting for minute-by-minute variations in irradiance, resulted in overestimations of daily stomatal conductance (13% mean error) and the magnitude of the error was directly related to the variation in irradiance for that day. We conclude that, provided variation in irradiance is accounted for, rates of stomatal opening and closing have little effect on estimates of daily water use.

Canopy transpiration and water fluxes in the xylem of the trunk of Larix and Picea trees — a comparison of xylem flow, porometer and cuvette measurements

Leaf gas exchange, transpiration, water potential and xylem water flow measurements were used in order to investigate the daily water balance of intact, naturally growing, adult Larix and Picea trees without major injury. The total daily water use of the tree was very similar when measured as xylem water flow at breast height or at the trunk top below the shade branches, or as canopy transpiration by a porometer or gas exchange chamber at different crown positions. The average canopy transpiration is about 12% lower than the transpiration of a single twig in the sun crown of Larix and Picea. Despite the similarity in daily total water flows there are larger differences in the actual daily course. Transpiration started 2 to 3 h earlier than the xylem water flow and decreased at noon before the maximum xylem water flow was reached, and stopped in the evening 2 to 3 h earlier than the water flow though the stem. The daily course of the xylem water flow was very similar at the trunk base and top below the lowest branches with shade needles. The difference in water efflux from the crown via transpiration and the water influx from the trunk is caused by the use of stored water. The specific capacitance of the crown wood was estimated to be 4.7 x 10-8 and 6.3 x 10-8 kg kg-1 Pa-1 and the total amount of available water storage was 17.8 and 8.7 kg, which is 24% and 14% of the total daily transpiration in Larix and Picea respectively. Very little water was used from the main tree trunk. With increasing transpiration and use of stored water from wood in the crown, the water potential in the foliage decreases. Plant water status recovers with the decrease of transpiration and the refilling of the water storage sites. The liquid flow conductance in the trunk was 0.45 x 10-9 and 0.36 x 10-9 mol m-2s-1 Pa-1 in Larix and Picea respectively. The role of stomata and their control by environmental and internal plant factors is discussed.