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.     

Effect of sowing date on the temperature response of leaf emergence and leaf size in barley

Abstract Rate of leaf emergence of barley grown in the field in each of 2 years was affected by sowing date and, where direct comparisons were possible, it was found that leaves on late-sown plants emerged more quickly. Rate of leaf emergence fluctuated throughout the season, slowing almost to zero in the winter. Much of this variation in rate was removed when the number of leaves was plotted against accumulated temperature rather than time. When emergence rates for each sowing were calculated using a common base temperature they were found to be well correlated with rate of change of daylength. However, it was (bund that base temperature as well as temperature response was affected by date of sowing. The pattern of change of size of leaves was also affected by date of sowing. It appeared that in low temperatures and short days, there was no increase in leaf size from leaf position to leaf position. The responses of leaf emergence, extension and final size to date of sowing appear to adapt the plant to grow quickly when sown early but to cease growth and possibly frost-harden at low temperatures.     

Analysis of leaf appearance,leaf death and phoma leaf spot,caused by Leptosphaeria maculans,on oilseed rape (Brassica napus) cultivars

Development of phoma leaf spot (caused by Leptosphaeria maculans) on winter oilseed rape (canola, Brassica napus) was assessed in two experiments at Rothamsted in successive years (2003–04 and 2004–05 growing seasons). Both experiments compared oilseed rape cultivars Eurol, Darmor, Canberra and Lipton, which differ in their resistance to L. maculans. Data were analysed to describe disease development in terms of increasing numbers of leaves affected over thermal time from sowing. The cultivars showed similar patterns of leaf spot development in the 2003–04 experiment when inoculum concentration was relatively low (up to 133 ascospores m⁻³ air), Darmor developing 5.3 diseased leaves per plant by 5 May 2004, Canberra 6.6, Eurol 6.8 and Lipton 7.5. Inoculum concentration was up to sevenfold greater in 2004–05, with Eurol and Darmor developing 2.4 diseased leaves per plant by 16 February 2005, whereas Lipton and Canberra developed 2.8 and 3.0 diseased leaves, respectively. Based on three defined periods of crop development, a piece-wise linear statistical model was applied to the progress of the leaf spot disease (cumulative diseased leaves) in relation to appearance (‘birth’) and death of leaves for individual plants of each cultivar. Estimates of the thermal time from sowing until appearance of the first leaf or death of the first leaf, the rate of increase in number of diseased leaves and the area under the disease progress line (AUDPL) for the first time period were made. In 2004–05, Canberra (1025 leaves ×°C days) and Lipton (879) had greater AUDPL values than Eurol (427) and Darmor (598). For Darmor and Lipton, the severity of leaf spotting could be related to the severity of stem canker at harvest. Eurol had less leaf spotting but severe stem canker, whereas Canberra had more leaf spotting but less severe canker.     

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.     

Environmental Influences on Panicle Differentiation and Spikelet Number of Pennisetum americanum

Pearl millet (Pennisetum americanum (L.) Leeke) has a juvenilephase after which the time to panicle initiation is reducedby short daylengths. To understand more fully the mechanismunderlying temperature ? daylength interactions on panicle initiationand differentiation, plants were grown (a) at a range of constanttemperatures under a short daylength from sowing until afterpanicle differentiation and (b) at one temperature until 20d after emergence and then at a range of temperatures duringa 10 d exposure to short daylength. Temperature prior to panicle initiation determined the numberof leaves initiated on the main stem and the size of the apicaldome at the start of panicle initiation. The number of leaves,in turn, influenced the duration of the phase from panicle initiationto anthesis: this phase required a constant thermal time whenexpressed as day degrees per leaf. At anthesis, panicle lengthwas positively correlated with the number of leaves on the mainstem (and temperature) prior to panicle initiation. Changingthe temperature only during exposure to inductive daylengthsaffected the rate of growth of the apical dome so that panicledifferentiation began within 10 d at high temperature (30?C)whereas differentiation did not commence in 10 dat 21?C. Paniclesdeveloped normally if differentiation had commenced under inductivedaylengths whereas panicles were abnormal when plants were returnedto long daylengths after panicle initiation but before visibledifferentiation. Relative extension rates of the panicle during differentiationwere correlated positively with temperature. The results areconsistent with the hypothesis that panicle initiation dependson the apex attaining a critical size and that temperature,by determining the number of leaves initiated on the main stem,affects the size of the apical dome and thus the onset of panicleinitiation, the duration of paniclc differentiation and thenumber of spikelets differentiated. Key words: Pennisetum americanum, panicle differentiation, spikelet number     

The Effects of Temperature and Daylength on the Rate of Development of Pigeonpea

Eleven cultivars of pigeonpea, representing five different maturitygroups, were grown under controlled conditions to determinehow the rate of development from sowing to flower bud initiation(FBI) and sowing to flowering was affected by temperature anddaylength, the two environmental variables thought to have mostinfluence on reproductive development. Both temperature and daylength had substantial effects overthe range 16–32 C and 10–14 h respectively. Comparisonswithin the 12 h daylength treatment showed no consistent patternwith mean temperature. A discrete-rate analysis was used todetermine the effect on development of each of the temperaturesmaking up the various treatment combinations of day and nighttemperature. The rate of development from sowing to FBI varied among cultivars.The responses to temperature were all strongly curvilinear withoptima between 20 and 24 C. The rate of development from sowingto flowering showed a similar pattern. The effect of daylengthon the rate of development was the greatest between sowing andFBI, with the greatest sensitivity between 12 and 14 h. For the range of conditions considered, temperature had at leastas great an influence as daylength on the rates of developmentfrom sowing to FBI and from sowing to flowering. Some of theresponses to time of planting that have previously been attributedto daylength effects may, in fact, have been due to temperature.The responses to temperature of reproductive and vegetativedevelopment are very different. Phenology, Cajanus cajan     

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.     

Growth and development of Avena fatua (Wild-oat) in the field

The growth and development of field-grown Avena fatua plants were studied for autumn and spring sowings in two consecutive years. The duration of various growth stages from sowing until anthesis was quantified in terms of thermal time (accumulated degree days) or photothermal time (degree days modified by photoperiod). Base temperatures and photoperiods for developmental phases were estimated as those which minimised the coefficient of variation among sowing dates. Relationships were derived between leaf emergence, canopy height, plant leaf area, and photothermal time. Stem extension and flowering occurred earlier in autumn-sown plants than spring-sown plants. Autumn-sown plants produced more leaves on the main stem, and had greater leaf area and above-ground biomass at anthesis than spring-sown plants.     

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 Influence of Temperature on Leaf Production and Vegetative Growth of Avena fatua

Leaf emergence, leaf area and plant d. wt of Avena fatua wereexamined under a range of constant day/night temperatures ata single light intensity under controlled environment conditions.A simple thermal time analysis was used as the basis for predictingplant growth and development. Multiple regression was used torelate leaf number to mean temperature and to the number ofdays from sowing to 50% plant emergence. From the thermal regressionequations a threshold temperature when growth ceases, the thresholdtime at which seedlings became established and emergence rateand a constant were also estimated. The reciprocals of the durationof leaf expansion and leaf number were linearly related; slopesand intercepts varied with temperature. Relationships were establishedbetween logarithms of leaf area, average temperature and daynumber. Final leaf area increased with average temperature andleaf number. Specific leaf area was inversely related to accumulatedtemperature. The logarithm of d. wt components, average temperatureand number of days from sowing were related; threshold temperature,relative growth rate and a constant were estimated. Parametervalues were compared and contrasted for the measures of growthand development. Shoot/root ratios were not clearly relatedto temperature. The consequences of the models are suggestedparticularly with respect to the formulation of a growth modelfor A. fatua Avena fatua L, wild oat, growth, development, temperature, thermal time     

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.     

Variability of phyllochron, plastochron and rate of increase in height in photoperiod-sensitive sorghum varieties

BACKGROUND AND AIMS: West African sorghum (Sorghum bicolor) varieties are generally highly photoperiod-sensitive, which is a necessary adaptation to the variable onset date of the rainy season and the variable dates of sowing in the savannah zone. Depending on sowing date, plants can produce from 12 to >40 leaves on the main culm, with height varying from 1 m to more than 5 m. The present study aimed to better understand the complex phenology of these variables. METHODS: A 2-year series of monthly sowings of three West African sorghum varieties was conducted near Bamako, Mali. Drought stress was avoided by supplemental irrigation. Rate of initiation of primordia at the stem apex was recorded, together with rate of leaf emergence and increase in plant height. KEY RESULTS: Leaf initiation and appearance rates (plastochron(-1) and phyllochron(-1)) were constant for a given sowing date in cases where less than 20 leaves were produced (generally observed with late sowing dates). In contrast, rates were bilinear for early sowing dates, for which plants produced more than 20 leaves. The secondary rates, which occurred from the 20th leaf onwards, were only half of the initial rate. Plastochron and phyllochron showed large variations among sowing dates, and were correlated with the rate of plant height increase. The initial plastochron and phyllochron were positively correlated with soil temperature and negatively correlated with both day length and day-to-day change of day length prevailing at plant emergence, but these factors explained only half of the variation observed. CONCLUSIONS: Although they belong to different genetic groups and have different height and photoperiod sensitivity, the three varieties studied exhibited similar response patterns of development rates among phenological phases and seasons, with the local landrace showing the greatest variation due to its longer vegetative phase and longer stem internodes. The possible adaptive advantages in African savannah environments of bilinear development rates and the associated limitation in height increase are discussed.     

Effects of temperature on the development and growth of winter wheat roots

Wheat plants were grown in columns of soil until early stem elongation at a wide range of constant root temperatures. Two light environments were imposed and three levels of nitrogen fertilizer added at sowing. Shoot and root development and growth were measured by destructive sampling to investigate the combined effects of temperature and changing nutrient and assimilate supply. Both mainstem leaf and root axis production were linearly related to thermal time above a base temperature of 0°C. Low irradiance affected the appearance of mainstem tillers and associated nodal root axes. Nitrogen had little effect on shoot or root development but increased shoot area between 6 and 8 mainstem leaves. Higher temperatures and supplementary light resulted in larger root systems when compared at equivalent times after sowing. Total root length and root dry weight increased exponentially with thermal time, based on the mean of 4 cm soil and 2 cm air temperatures, but no single relation existed for all temperature and light treatments. Total plant dry matter, root length and root dry weight increased linearly with accumulated, intercepted, photosynthetically active radiation. Root growth responded less than the shoot to supplementary light. Increasing temperature reduced the proportion of root weight to total plant weight.     

Responses of potato genotypes to drought. I. Expansion of individual leaves and osmotic adjustment

The expansion of individual leaves was examined in 19 genotypes of potato (Sofunum tuberosum L.) grown either with irrigation or droughted from the time of plant emergence. Drought reduced the thermal time from emergence to leaf appearance in 10 genotypes but had no significant effect in the other genotypes. In all the genotypes, final size of leaves was reduced by drought, but the magnitude of the effect differed significantly (P < 0.001) with genotype. In the droughted treatment, the final size of leaves was correlated (P < 0.001) with the maximum rate of leaf expansion suggesting that reductions in final size of leaves were the result of reduced expansion rate rather than of effects on the duration of expansion. Both the constant and the slope of the relation between leaf expansion rate and soil moisture deficit differed between genotypes. Osmotic adjustment was limited, maximally 0.16 MPa, and did not correlate with the ability to maintain leaf expansion with increasing soil moisture deficit.     

Leaf growth and demography of the rheophytic fern Thelypteris angustifolia (Willdenow) Proctor in a Puerto Rican rainforest

Morphological and temporal aspects of the growth of leaves of Thelypteris angustifolia (Willdenow) Proctor are under study along a 400-m section of the Sonadora River at the El Verde Field Station in the Luquillo Experimental Forest of Puerto Rico. Emergence, expansion and growth of leaves of 149 sporophytes have been observed approximately mid-month every January, May and September since September 1991. Results of the first four years of this long-term study of T. angustifolia are reported here. Although they are land plants, sporophytes of T. angustifolia are rheophytic, subject to intermittent flooding and submerged approximately 7% of the time. The pinnate leaves of T. angustifolia exhibit sterile-fertile leaf dimorphy. Only 7.6% of the leaves in the the sample were fertile. The petiole of a fertile leaf was 41% longer than that of a sterile leaf while leaf blades were the same length. Leaves expanded at a mean rate of 1.7 cm per day, maturing in 29.7 days. Mean leaf life span was 10.8 months with a maximum of 26 months. Sterile leaf longevity (11.0 months) was longer than that of the fertile leaf (9.6 months). Leaves were produced at a mean rate of 4.7 leaves per plant per year. Leaf damage occurred on 38% of the leaves observed. Net leaf counts for individual sporophytes ranged from one to eight with a mean of 3.0 leaves per plant. Fertile plants had higher mean leaf counts (4.1) than plants without fertile leaves (2.8). Leaf production, although possibly triggered by increasing daylength, also paralleled seasonal increases in rainfall during the wetter mid-May/September months. Increased leaf mortality occurred during the drier mid-January/May period.     

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.     

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.     

Regulation of photosynthate partitioning into starch in soybean leaves : response to natural daylight

Studies conducted in controlled environments indicate that daylength affects the proportion of photosynthate stored in leaves as starch or sucrose. To examine the response of partitioning to natural daylight, soybeans (Glycine max [L.] Merr. cv Williams) were grown at 12 different times between May and November in a constant temperature greenhouse without supplemental lighting. Plants were transferred from the greenhouse to a controlled environment chamber at the end of civil twilight at a set developmental stage (expanding seventh trifoliolate leaf, counting acropetally). Net photosynthesis and the accumulation of starch and sugar in fully expanded fourth trifoliolate leaves were determined the following day under standard conditions in the chamber (lights-on synchronized with sunrise). Photosynthesis on a leaf area basis decreased about 10% between midsummer and early autumn. Leaf soluble sugar accumulation was low at all harvests. However, a twofold increase in photosynthate partitioning into starch occurred over the same time period, resulting in an 80% increase in absolute starch accumulation rate. Starch was responsible for about 78% of the increase in leaf dry matter during the light at all harvests, indicating that starch accumulation as affected by prior daylight conditions will alter export of photosynthate during the light period. Photosynthate partitioning into starch was linearly correlated with daylength at harvest, prior average peak solar irradiance, and other parameters that correlated with daylength and solar radiation such as harvested top dry matter. The relation between growth and seasonal changes in daylight (including daylength, irradiance, and light integral) are discussed in relation to photosynthate partitioning under field conditions.     

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.