Field studies of leaf development and expansion in the leek (Allium porrum)

An analysis of leek leaf development and expansion was carried out over three seasons using field-grown plants of three varieties which were directly sown at different dates or transplanted from controlled conditions. In all cases, successive leaves appeared (tip visible) at equal intervals of accumulated temperature. Detailed analysis of a single sowing in 1985 showed that the regularity of leaf appearance was a consequence of the coordinated response to accumulated temperature of leaf initiation (plastochron 100°C days > 0°C) and leaf blade and sheath extension. For each successive leaf, an additional 32°C days were required between initiation and appearance to allow for the linear increase in ‘sheath’ height, giving a phyllochron of 132°C days. Direct measurement of leaf extension before and after leaf appearance, and of the length of the leaf extension zone, confirmed that the rate of leaf extension, in terms of accumulated tempeature, was constant, and independent of leaf number. However, there were differences between seasons and between varieties in the responses of leaf appearance, leaf extension and ‘sheath’ length to accumulated temperature. It was concluded that the simple ontogenetic increase in leaf dimensions, which was a feature of all the crops studied, was a consequence of the progressive increase in the duration of leaf expansion.     

The influence of temperature on leaf development and growth in potatoes in controlled environments

Three controlled environment experiments were conducted at different temperatures to determine the relation between temperature and leaf development and growth in the potato (cv. Maris Piper). Developmental stages are defined for the appearance and duration of leaf extension in the potato and comparisons made with other temperate zone crops. The rate of leaf appearance was linear over the temperature range (9–25°C) and above 25°C there was no further increase in the rate. The temperature coefficient for the rate of appearance of leaves was 0.032 leaves (degree days)^-1 using a base temperature of 0°C. The duration of extension of an individual leaf decreased with increase in temperature up to 25°C such that the thermal duration was constant at 170 degree days using a base temperature of 0°C for leaf positions 4–10 on the main stem. At higher leaf positions the thermal duration was similar or greater. The advantages and limitations of controlled environment work as a parallel to field experimentation are discussed.     

The Effects of Position and Temperature on the Expansion of Leaves of Vicia faba L.

The growth in area of the first eight leaves of broad bean plantswas investigated in growth room experiments. Plants were grownat either 20 or 14 °C or transferred from 20 to 14 °C.Rates of leaf appearance and unfolding increased with temperature.The duration of growth of a leaf increased with leaf numberfor the first five leaves and then remained constant The meangrowth rate declined or remained constant with increasing leafnumber Durations of growth were shorter and growth rates largerat 20 °C than at 14 °C Plants responded immediatelyto the change in temperature Final areas of leaves which expandedafter transfer from 20 to 14 °C were larger than those grownat 20 °C Vicla faba L., broad bean, leaf expansion, temperature responses     

Temperature Effects on Phenological Development and Yield of Muskmelon

Our goal was to construct a simple muskmelon phenology modelthat could be run with easily obtainable weather station dataand used by growers to quantify phenological development andaid in projecting harvest dates. A growth chamber experimentwas conducted with two cultivars of muskmelon (‘Gold Rush’and ‘Mission’) to determine how main vine leaf appearancerates responded to temperature. We identified three cardinaltemperatures for leaf appearance rate: the base temperature(10 °C) at which leaf appearance rate was zero; an optimumtemperature (34 °C) at which the rate of leaf appearancewas maximal; and an upper threshold temperature (45 °C)at which leaf appearance rate returned to zero. Using thesethree cardinal temperatures, we constructed a simplified thermalunit accumulator for hourly measurements of air temperature.Main vine plastochron interval (PI), thermal time to harvest,and final yield were determined for three cultivars of muskmelon(‘Explorer’, ‘Gold Rush’ and ‘Mission’)grown in the field at Overton, TX, USA, over six transplantingdates from March to June 1998. PI was calculated for each cultivarx transplanting date combination as the reciprocal of the slopeof main vine node number vs. accumulated hourly thermal units(     

Variation in growth attributes of contrasting populations of Trifolium repens

A range of growth attributes was measured in seedlings of 10 Trifolium repens populations, differing in leaf size and origin, grown in three temperature and two glasshouse environments. Growth rates of large leaf types of Mediterranean origin were higher than those of smaller leaf types at 10°C. However, the greater temperature response of the smaller leaf types resulted in higher growth rates for S.100 and S.184 than for a large leaf type from Israel at 20°C. The increase of growth rate with temperature was associated with changes in leaf area ratio and net assimilation rate between 10° and 15°C but only with changes in net assimilation rate between 15° and 20°C. Within each temperature environment, population differences in growth rate were related to differences in net assimilation rate rather than leaf expansion. At low temperature a greater proportion of dry matter was distributed to leaf tissue in large leaf types particularly those of Mediterranean origin but they showed a proportionately smaller increase in allocation to leaves with increasing temperature compared with small leaf types. In the glasshouse environments growth rates in spring were more than double those in the autumn. This difference was associated with net assimilation rates which were about five times greater in the spring environment. However, leaf area ratios in the spring were only half those in the autumn. These differences in leaf area ratio between the glasshouse environments were closely related to differences in specific leaf area but not to differences in distribution of dry matter to leaf tissue which was greater in the spring environment.     

Is thermal time adequate for expressing the effects of temperature on sunflower leaf development?

We have tested whether the effects of temperature on sunflower leaf growth could be documented by using thermal time. The rates of leaf expansion and of cell division were analysed in leaves located at two positions on the stem, and a spatial analysis of expansion rate was carried out. Experiments were performed in growth chamber (stable conditions), in the field or in a greenhouse (fluctuating conditions). We compared three methods for characterizing the rate and the duration of expansion. Responses to leaf temperature were consistent only when expansion was characterized as a two-phase process — a period of exponential expansion (constant relative expansion rate, RER ) followed by a decrease in RER . RER and relative cell division rate ( RDR ) responded linearly to temperature with a common response curve for all studied conditions. This response curve was also common to all studied zones within a leaf and to leaves at two positions on the stem. The reciprocals of the durations of the periods of exponential expansion, non-zero expansion and non-zero division were also linearly related to leaf temperature with common response curves in a given leaf zone. The x -intercepts of all these response curves and of the response curve of leaf initiation rate to temperature did not significantly differ in an analysis of covariance, with a common value of 4·8 °C. The expression of time in cumulative degree days, with a base temperature of 4·8 °C, resulted in a unique time course of RER and cell division rate regardless of temperature. These results suggest that a powerful 'program' of leaf development exists in a sunflower plant.     

Effect of temperature, irradiance and photoperiod on morphological characters of seedlings of contrasting white clover populations

Seedlings of 10 contrasting white clover populations differing in leaf size and origin were grown in a glasshouse in spring and autumn, and in controlled environments in which temperature (10°, 15°, 20°C), photoperiod ( 8 , 16-h) and irradiance ( 13, 47 , 57 Wm^-2) were varied. There were large effects of these environmental variables on a range of morphological characters. Population differences were closely related to classification of the populations by leaf size. Within the large leaf types there was also an effect of origin. In particular, Mediterranean types differed from those of northern origin. Rate of leaf appearance increased with temperature but was less affected by photoperiod. Population differences were greater at lower temperatures. Petiole length was more sensitive to temperature than was leaflet length, particularly in the range 10° to 15°C. Similarly, petiole length increased appreciably with photoperiod extension; smaller leaf types responded more than large leaf types with the result that there were no population differences in the 16-h photoperiod. Population differences in the other environments were closely related to leaf size classification. The onset of stolon production was delayed by lower temperatures and by short photoperiods but stolon number at the sixth leaf stage of development was not closely related to this character in the temperature environments since stolon numbers increased at lower temperatures. In low temperature and short day conditions the large leaf types including those of Mediterranean origin showed a desirable combination of long petioles and large leaves compared with the smaller leaf types. However, these differences between the two groups were eliminated for petiole length, and much reduced for leaflet length, by the greater response to temperature and photoperiod of the smaller leaf types. The implications of these responses for seasonal production are discussed. There were large effects of irradiance on the lengths of leaflets and petioles with longer ones at higher irradiance levels. There were also faster rates of leaf appearance at higher light levels. Stolon production was faster at 57 Wm^-2 than at 47 Wm^-2 while at 13 Wm^-2 no stolons were produced. After an extended period in the lowest light environment there was some plant mortality.     

The effects of temperature on leaf growth of sugar beet varieties

Leaf growth of nine varieties of sugar beet (Beta vulgaris L.) was studied at constant temperatures of 7, 11, 15 and 20·C, using generalised logistic curves fitted to the data to estimate the parameters of growth. The rate of leaf appearance increased linearly with temperature and was the same in all varieties. There were differences between varieties in the weighted mean rates of expansion of leaf area per plant (ā), the temperature coefficient of ā and the leaf area duration (D); these differences were caused more by differences in rates of expansion and final sizes of individual leaves than by differences in rates of leaf production. The growth of the first six leaves produced by each plant was examined in detail. The greater size of successive leaves of plants and genotypic differences between comparable leaves were more attributable to differences in the rate than differences in the duration of leaf expansion. Increasing temperatures increased leaf size because they accelerated the rate of expansion more than they shortened the duration of the expansion phase. It is inferred that all effects arose through differences in the initial sizes of leaves before they unrolled from the shoot apex. Dry matter production was proportional to D but was partitioned more to the storage root at the colder temperatures. This may have been related to the differential effects of temperature on cell division and expansion and the relative contribution of these two processes to the final sizes of the leaves and storage root.     

The effect of temperature on leaf appearance and canopy establishment in fibre hemp (Cannabis sativa L.)

The effects of temperature on the development and growth of hemp (Cannabis sativa L.) have never been quantified. Therefore, to establish the effect of temperature on leaf appearance and canopy establishment of fibre hemp under controlled and field conditions, plants were grown in growth chambers at 11 regimes with average temperatures between 10°C and 28°C, and three cultivars were sown in the field in March, April and May in 1990, 1991 and 1992. In the field, thermal time (base 0°C) between sowing and emergence ranged from 68°Cd to 109.5°Cd (average 88.3°Cd). Rates of leaf appearance and stem elongation increased linearly with temperature between 10°C and 28°C. The base temperature for leaf appearance was 5.7°C from the growth chamber experiments and 1°C from the field experiments. In the field, the base temperature for the relationship between light interception by the canopy and thermal time was 2.5°C, and thermal time, calculated at the appropriate base temperature, accounted for about 98% of the variance in the number of leaves and for 98.6% of the variance in the proportion of light intercepted by the canopy. Days from emergence accounted for less of the variance in both parameters than thermal time. Interception of 90% of light was attained on average at 465°Cd (base 0°C) after emergence. It is concluded that thermal time is a simple and accurate tool to describe leaf appearance and light interception in fibre hemp.     

Variation in growth of overwintered stolons of contrasting white clover populations in response to temperature, photoperiod and spring environment

The response of overwintered stolons of nine contrasting white clover populations to temperature, photoperiod and natural conditions was studied in six environments during the spring. Rate of leaf appearance, leaflet length, petiole length, stolon internode length, dry matter distribution within the plant and total dry weight were measured on 15 plants of each population/environment combination. Most characters, except leaf size and proportion of dry matter allocated to leaf, responded positively to temperature in the range 10 – 20°C. A positive effect of photoperiod extension was also found for all characters except rate of leaf appearance, internode length and distribution of dry matter to leaf. Population differences in response to environment were found which were related to both leaf size classification and origin. Stolon dry weight was positively correlated with leaf length, petiole length and stolon internode length in most environments. The relationships between the eight characters were often complex and canonical variate analysis provided a convenient way to discriminate between the populations based on all eight characters.     

Field Studies of Cereal Leaf Growth: I. INITIATION AND EXPANSION IN RELATION TO TEMPERATURE AND ONTOGENY

Measurements of leaf initiation, appearance, and expansion arepresented for winter wheat and spring barley crops. For winterwheat, these processes occurred during periods of several weekswhen fluctuating temperatures influenced process rates. Analysisof these measurements was facilitated by plotting variablesagainst the time integral of temperature above an appropriatebase temperature (O °C), here called thermal time with unitsof °C d. Leaf primordial number and appearance stage increasedlinearly with thermal time for both winter wheat and springbarley which initiated 12 and 9 leaves respectively. When plottedagainst thermal time 90% of laminar and leaf length growth and80% of laminar width growth was satisfactorily described bya straight line for both species. This enabled an average extensionrate and duration of linear growth to be defined for each leaf.When expressed in thermal time, wheat leaves had a similar durationof linear growth (210 °C d; s.d. 30 °C d) with insolationexerting a negligible influence. The first seven barley leaveshad a shorter duration of linear growth (151 °C d; s.d.8 °C d). For wheat, final leaf length and laminar widthincreased with leaf number and were not apparently associatedwith changes in apical development stage. Changes of barleyleaf dimensions with leaf number were more complex.     

The setting of rates of development of wheat plants at crop emergence: Influence of the environment on rates of leaf appearance

Examination of the time courses, expressed in terms of accumulated temperature over 0°C, of leaf appearance of fifteen sowings of winter wheat, including five cultivars, over three seasons at Auchincruive, revealed several examples of bi-linear rather than linear responses. Stepwise linear regression analysis, varying the temperature threshold by intervals of 1°C, showed that a linear relationship could be obtained for each sowing, but the appropriateness of this analysis is questioned. An alternative interpretation, that such changes in rate do have biological rather than purely statistical significance, is supported by the fact that, when they occurred, they coincided with the change in rate of spikelet initiation by the stem apex. Furthermore, similar relationships between leaf appearance and reproductive development have been demonstrated for perennial ryegrasses. Analysis of each sowing also revealed the existence of sub-populations of plants with different rates of leaf appearance and final leaf numbers, but with synchronous phenologies. These findings are discussed in relation to the coordination of the development of apex and leaves, and the environmental cues required to ensure this coordination.     

The prediction of leaf canopy expansion in the leek from a simple model dependent on primordial development

In an analysis of leaf development of leek plants grown in the field in 1988, successive leaves initiated, appeared (tip and ligule) and senesced at equal intervals of accumulated temperature/thermal time. These intervals corresponded to a plastochron of 92°C days and phyllochrons of 135 (tip) and 233 (ligule) °C days. The rate of appearance of ligules was exactly equal to the rate of leaf senescence, with the result that the number of fully-expanded leaves per plant remained constant at 1.4. These data, which were compatible with results from previous seasons, were used to develop a model of the interrelationships between primordium initiation at the shoot apex and subsequent events in the development of individual leaves. Primordium initiation is considered to be the primary controlling event in the life of a leaf, and the processes of tip appearance, ligule appearance and death can be predicted from knowledge of the number of primordia which have been initiated, without reference to the environment. A model of canopy expansion, based on the central role of the shoot apex, was developed using the temperature relations of primordium initiation and additional data on leaf expansion and leaf dimensions. Leaf area indices computed in this way provided a satisfactory simulation of the thermal-time course of leaf area index observed in a previous season, 1985.     

Diurnal variation in the temperature response of leaf extension of two bunchgrass species in the field

The effect of temperature on short-term leaf extension rates was studied for two cool-season tussok grasses, Agropyron desertorum and Pseudoroegneria spicata, growing in the field under a variety of water stress and defoliation conditions. Leaf extension rates and air temperatures were monitored every half hour during numerous 12- to 65-h periods in three growing seasons using auxanometers constructed of precision resistors. For both species, a three-phase relationship between leaf extension rate and temperature was observed during diurnal cycles. Leaf extension rate increased linearly with temperature from dawn until midday (phase 1). Leaf extension then increased rapidly, reaching maximum rates in the early evening (approximately 1900h), despite decreasing temperatures during this period (phase 2). Finally, leaf extension rate declined with temperature from evening until dawn (phase 3). This diurnal cycle was described by linear (phase 1) and quadratic (phases 2 and 3 combined) regression models. Although the rate of leaf extension and daily integrals were affected by the water stress and defoliation treatments, the diurnal pattern was consistently observed. Temperature was probably a major factor governing leaf extension rates at night (phase 3), but it appeared unimportant in controlling leaf extension between dawn and midday. The relative importance of physiological and environmental factors controlling leaf extension rate appears to shift during the day in these species under field conditions.     

Leaf Extension in Zea mays: II. LEAF EXTENSION IN RESPONSE TO INDEPENDENT VARIATION OF THE TEMPERATURE OF THE APICAL MERISTEM, OF THE AIR AROUND THE LEAVES, AND OF THE ROOT-ZONE

The temperatures of the roots, the apical meristem, and theshoots of Zea mays plants were varied independently of eachother and the rates of leaf extension were measured. When thetemperature of the apical meristem and region of cell expansionat the base of the leaf was kept at 25 °C, changes of leafextension in response to changes of root and shoot temperatureswere less pronounced. When the temperature of the meristematicregion was changed by increments of 5 or 10 °C from 0 to40 °C, and the root and shoot temperatures were kept at25 °C, rapid changes in leaf extension occurred. It was concluded that the rates of leaf extension were controlledat root-zone temperatures of 5 to 35 °C by heating or coolingof the meristematic region. Changes in rates of leaf extensionin response to changes in air temperature were attributed todirect effects on the temperature of the meristematic regionand on the physiology of the leaf.     

Spring Wheat Leaf Appearance and Temperature: Extending the Paradigm?

Extensive research shows temperature to be the primary environmental factor controlling the phyllochron, or rate of leaf appearance, of wheat (Triticum aestivum L.). Experimental results suggest that soil temperature at crown depth, rather than air temperature above the canopy, would better predict wheat leaf appearance rates. To test this hypothesis, leaf appearance in spring wheat ('Nordic') was measured in a 2-year field experiment (Nunn clay loam soil; fine, smectitic, mesic Aridic, Argiustoll) with three planting dates and two soil temperature treatments. One temperature treatment (denoted +3C) consisted of heating the soil at crown depth to 3 degrees C above the ambient soil temperature (denoted +0C). Main stem cumulative leaf number was measured at least weekly until flag leaf emergence. Leaf appearance was essentially linear with both air and soil growing degree-days (GDD), although there was a stronger linear relationship with soil GDD in the +0C plants than in +3C plants. A weak positive relationship between planting date and the phyllochron was observed. Unexpectedly, we found that heating the soil did not increase the rate of leaf appearance, as the paradigm would predict. To explain these results, we propose extending the paradigm in two ways. First, three processes are involved in leaf appearance: (1) cell division at the shoot apex forms the primordium; (2) cell division in the intercalary meristem forms the cells that then (3) expand to produce the leaf. Cell division is predominantly controlled by temperature, but cell expansion is considerably more affected by factors other than temperature, explaining the influence of other factors on the phyllochron. Secondly, the vertical distribution of the two meristems and region of cell expansion occur over a significant distance, where temperature varies considerably, and temperature at a specific point (e.g. crown depth) does not account for the entire temperature regime under which leaves are developing.     

Daylength change and leaf appearance in winter wheat

Abstract In the field successive leaves of winter wheat appear at a rate which varies because it depends strongly upon temperature. When plotted against ‘thermal time’, however (temperature accumulated above a fixed base of 0°C), leaf appearance was a strictly linear function of temperature. The mean rate of leaf appearance in thermal time, R′, was faster for a spring sowing than for an autumn sowing. The variation in R′ between sowings was better correlated with the rate at which daylength was changing when the plants emerged than with the mean daylength while leaves were appearing.     

Temperature dependence of growth,development, and photosynthesis in maize under elevated CO2

Global atmospheric carbon dioxide concentrations (C_a) are rising. As a consequence, recent climate models have projected that global surface air temperature may increase 1.4–5.8 °C with the doubling of C_a by the end of the century. Because, changes in C_a and temperature are likely to occur concomitantly, it is important to evaluate how the temperature dependence of key physiological processes are affected by rising C_a in major crop plants including maize (Zea mays L.), a globally important grain crop with C₄ photosynthetic pathway. We investigated the temperature responses of photosynthesis, growth, and development of maize plants grown at five temperature regimes ranging from 19/13 to 38.5/32.5 °C under current (370 μmol mol⁻¹) and doubled (750 μmol mol⁻¹) C_a throughout the vegetative stages using sunlit controlled environmental chambers in order to test if the temperature dependence of these processes was altered by elevated C_a. Leaf and canopy photosynthetic rates, C₄ enzyme activities, leaf appearance rates, above ground biomass accumulation and leaf area were measured. We then applied temperature response functions (e.g., Arrhenius and Beta distribution models) to fit the measured data in order to provide parameter estimates of the temperature dependence for modeling photosynthesis and development at current and elevated C_a in maize. Biomass, leaf area, leaf appearance rate, and photosynthesis measured at growth C_a was not changed in response to CO₂ enrichment. Carboxylation efficiency and the activities of C₄ enzymes were reduced with CO₂ enrichment indicating possible photosynthetic acclimation of the C₄ cycle. All measured parameters responded to growth temperatures. Leaf appearance rate and leaf photosynthesis showed curvilinear response with optimal temperatures near 32 and 34 °C, respectively. Total above ground biomass and leaf area were negatively correlated with growth temperature. The dependence of leaf appearance rate, biomass, leaf area, leaf and canopy photosynthesis, and C₄ enzyme activities on growth temperatures was comparable between current and elevated C_a. The results of this study suggest that the temperature effects on growth, development, and photosynthesis may remain unchanged in elevated C_a compared with current C_a in maize.     

Nutrient uptake by field-grown broccoli and net nutrient mobilization during inflorescence development

Our objective was to determine the effect of temperature on root appearance at successive stalk nodes of a maize (Zea mays L.) plant. Plants were grown in three controlled temperature regimes (30/24, 26/20 and 19/14°C 16-/8-h day/night or 28, 24 and 17.3°C mean), as well as in the field. Three plants were dissected twice a week for morphological observation. A second-degree polynomial, with time as the independent variable and with coefficients linearly related to temperature, was fitted to the controlled-environment data. This equation tested well against two sets of field data. The appearance of newly rooted stalk nodes was related to the appearance of new leaf tips and collars. Such information is urgently needed for predicting leaf, stalk and root growth in maize and other grasses.     

Influence of day length and temperature on number of main stem leaves and time to flowering in lupin

A growth chamber experiment was carried out to investigate the influence of day length and temperature on the development of flowering in eight varieties of the three grain lupin species Lupinus albus (Wat and C3396), L. angustifolius (Gungurru, Polonez and W26) and L. luteus, (Juno, Radames and Teo). The plants were grown at two temperatures, 10°C and 18°C, in combination with five daylength regimes: 10, 14, 18, 24 h day at full light intensity and 10 h full light extended with 8 h low intensity light. Increased daylength decreased days from sowing to flowering in all varieties, but had little effect on thermal time to flowering in most varieties. However, C3396, W26 and Radames had a significantly longer thermal time to flowering at high, non‐vernalising temperature (18°C) at short daylengths. Low light intensity daylength extension did not significantly influence thermal time to flowering. For flower initiation, measured as number of leaves on the main stem three types of response were found. All varieties formed fewer leaves on the main stem at 10°C than at 18°C, although the two thermo‐neutral varieties of L. luteus, Juno and Teo, gave only a small response to temperature and daylength. In Polonez, Gungurru and Wat, low temperature decreased leaf number, but there was only a small response to changes in daylength. Three varieties, C3396, W26 and Radames, showed longer thermal time to flowering at 18°C with short daylengths. This could be explained by a greater number of main stem leaves formed at short daylength at non‐vernalising temperatures. Increased daylength decreased leaf number in these varieties, but never to a smaller number than for plants grown at 10°C. In these varieties, low intensity extension of the daylength had a similar (W26, Radames) or decreased (C3396) effect compared to full light extension. The hastening of time to flowering by long days could be separated into two effects: a high light energy effect hastened development by increasing the rate of leaf appearance in all varieties, while low light energy in thermo‐sensitive varieties was able to substitute for vernalisation by decreasing leaf number.