Temperature and daylength control of flower initiation in winter oilseed rape (Brassica napus L.)

The influence of low temperature and daylength on pre-floral growth and flower initiation in winter oilseed rape cv. Mikado was examined under controlled environment conditions at the University of Newcastle upon Tyne during 1985 and 1986.
The vernalisation requirement of Mikado was most effectively fulfilled by temperatures of 6 °C and 9 °C. Plants maintained at both higher and lower temperatures had an extended pre-floral growth phase. The transition from vegetative to reproductive growth in plants maintained at 12 °C was delayed by slow accumulation of the cold requirement, whereas flower initiation appeared to be delayed by limited leaf production, dry matter accumulation and/or assimilate availability in plants grown at 3 °C. The mechanism of floral induction remained unresolved but it was clear that flower initiation was not controlled by low temperature per se . Short days partially substituted for the cold requirement at 12 °C but photoperiodic induction of flower initiation was less important than the influence of low temperature.     

Determinism of carbon and nitrogen reserve accumulation in legume seeds

In legume plants, the determination of individual seed weight is a complex phenomenon that depends on two main factors. The first one corresponds to the number of cotyledon cells, which determines the potential seed weight as the cotyledon cell number is related to seed growth rate during seed filling. Since cell divisions take place between flowering and the beginning of seed filling, any stress occurring before the beginning of seed filling can affect individual seed growth rate (C and N reserve accumulation in seeds), and thus individual seed weights. The second factor concerns carbon and nitrogen supply to the growing seed to support reserve accumulation. Grain legume species produce protein-rich seeds involving high requirement of nitrogen. Since seed growth rate as determined by cotyledon cell number is hardly affected by photoassimilate availability during the filling period, a reduction of photosynthetic activity caused by nitrogen remobilization in leaves (e.g., remobilization of essential proteins involved in photosynthesis) can lead to shorten the duration of the filling period, and by that can provoke a limitation of individual seed weights. Accordingly, any biotic or abiotic stress during seed filling causing a decrease in photosynthetic activity should lead to a reduction of the duration of seed filling.     

Effects of below- and above-ground herbivores on plant growth,flower visitation and seed set

Separate and combined effects of root and leaf herbivores on plant growth, flower visitation and seed set were tested in a factorial experiment using potted mustard, Sinapis arvensis, at an old fallow field. A 50% leaf removal by cabbageworms (Pieris rapae) when the seedlings had their first four leaves reduced plant height and shoot mass, and delayed the onset of flowering. Root herbivory by two wireworms (Agriotes sp.) over the whole experiment changed flower visitation; the number of flower visitors per plant was higher in plants with root herbivores than in plants without root herbivores. Combined leaf and root herbivory affected flowering period, number of fruits per plant and number of seeds per fruit. Plants attacked by leaf and root herbivores had a shorter flowering period and produced fewer fruits per plant than plants with root herbivores only. Although the experimental plants faced major herbivore-induced growth changes, plant reproduction (seed set and weight per plant) was similar in all treatments, documenting their ability to effectively compensate for leaf and root herbivory.     

Description and Model of the Architecture of Four Genotypes of Determinate Autumn-sown White Lupin (Lupinus albus L.) as Influenced by Location, Sowing Date and Density

The structure of four new autumn-sown determine genotypes ofwhite lupin, grown with a combination of sowing dates and plantdensities at four locations in France was studied in 1990/91and 1991/92. Plant architecture was characterized in terms ofthe number of first-order branches produced and the distributionof leaves on these branches and how they varied with the numberof mainstem leaves and the position of the branch on the mainstem. The variation in the number of mainstem leaves on determinegenotypes was satisfactorily described by an earlier model developedfor indeterminate genotypes using the intensity of vernalizationrelative to accumulated temperature above 3°C as input.However the architecture of the first-order branches, definedby their number and the total number of leaves per branch ateach subtending position on the mainstem, differed and was influencedby genotype, location and sowing date. In four determinate genotypes,these two characters were positively correlated, and correlatedwith the number of leaves on the mainstem. Increasing densityfrom 10 to 40 plants m^-2 reduced the number of branches andtheir total number of leaves. Pruning 15 leaves from the mainstemat the beginning of stem elongation did not influence the developmentof the first-order branches. There appeared to be a functionalrelationship between the number of leaves on the mainstem andthe number of branches established very early in the growingseason. The profiles of number of leaves on the first-order brancheson the mainstem with branch position were modelled using twosub-models, one describing the mean number of leaves on thebranches actually present, and the second describing the frequentlyof occurrence of a branch at a particular position on the mainstem.Of the five model parameters, one, representing the minimalnumber of branches present on the plant, was correlated withthe number of first-order branches. Genotype, location, sowingdate and density affected the shape of the profiles.Copyright1993, 1999 Academic Press While lupin, Lupinus albus L., determine growth, architecture, model, branch     

Studies on the Photoperiodic Responses in Short-Day Plant Helianthus tubevotus

Plants of Helianthus tuberosus, variety white tuber, were treated with various daylengths of 4, 6, 8, 10, 11, 12, 13, 14, 15, 18 h. for 25 days as soon as six leaves on a plant appeard Irradiation for 6–13 h per day induced the plants to form flower buds and flowering, daylength with 14 h or longer kept the plants in vegetative growth. The experiments showed that this variety of Helianthus tuberosus required short days for flowering and the critical daylength was about 13 h. The plants were treated with short days for different durations. At least 17 days were required, Formation of flower buds and flowering had positive correlation with the number of short days over 17 days. After short-day induction, the shorter the daylength is, the more the flower buds inverted. Long-day treatment after an appropriate period of short days wouid reduce the number of flower and induce new vegetative branches from flowering granehes.     

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     

Acclimation to High CO(2) in Monoecious Cucumbers : I. Vegetative and Reproductive Growth

CO₂ concentrations of 1000 compared to 350 microliters per liter in controlled environment chambers did not increase total fruit weight or number in a monoecious cucumber (Cucumis sativus L. cv Chipper) nor did it increase biomass, leaf area, or relative growth rates beyond the first 16 days after seeding. Average fruit weight was slightly, but not significantly greater in the 1000 microliters per liter CO₂ treatment because fruit numbers were changed more than total weight. Plants grown at 1000 and 350 microliters per liter CO₂ were similar in distribution of dry matter and leaf area between mainstem, axillary, and subaxillary branches. Early flower production was greater in 1000 microliters per liter plants. Subsequent flower numbers were either lower in enriched plants or similar in the two treatments, except for the harvest at fruiting when enriched plants produced many more male flowers than the 350 microliters per liter treatments.     

COMPONENTS OF SEED AND POLLEN YIELD OF LOBELIA CARDINALIS: VARIATION AND CORRELATIONS

Plants from three Lobelia cardinalis populations were grown under common garden conditions to assess intra- and interplant variation in seed and pollen production. Seed number per flower and mean seed weight varied systematically with floral position on the inflorescence (lowest values were from terminal flowers) but pollen grain number per flower did not vary systematically with floral position. Most of the remaining variance in seed and pollen grain number per flower and mean seed weight was distributed among plants; clones produced very similar amounts of pollen and seed. Seed yield was positively correlated with seed production per flower and with total flower production, but not with mean seed weight; pollen yield was also positively correlated with pollen grain production per flower and total flower production. Seed and pollen yield were simple linear functions of plant size but only pollen yield was a simple linear function of flower production; seed yield was a quadratic function in which the second order term was negative. This quadratic relationship resulted from a negative correlation between seed number per flower and total flower production. This correlation, in addition to the wide variation among plants in pollen number per flower, accounts for the weak correlation of seed and pollen yield. I conclude from these data that it is unlikely that plants in natural L. cardinalis populations transmit genes to the population's seed crop equally through pollen and ovules—emphasizing the importance of measuring both male and female components of reproductive success.     

The response of sorghum and sunflower to short-term waterlogging

Summary The effect of waterlogging on sunflower and sorghum was investigated in relation to stage of development (sunflower-6-leaf, buds-visible, anthesis; sorghum-5-leaf, initiation, anthesis) and duration of waterlogging (3, 6 and 9 days) under glasshouse conditions. Additionally, the potential adaptation of the two crops was observed by waterlogging some plants at all three growth stages. With sunflower, leaf expansion and stem extension were inhibited by waterlogging at the 6-leaf and buds-visible stage although these effects did not always persist until maturity while, with anthesis waterlogging, rapid desiccation of leaves was observed. Yield was most affected by the anthesis waterlogging but no consistent effect on seed number or 1000 seed weight was recorded.Waterlogging sorghum plants suppressed normal tillering but had little effect on dry weight of the main stem. Late tillering was stimulated by waterlogging. Reductions in leaf area occurred at all stages of development in response to waterlogging with these effects being more marked at initiation. Similarly, yield was most reduced by the initiation waterlogging largely as a result of reduced seed number.In neither species was there a clear relationship between duration of waterlogging and subsequent reduction in growth and yield. With respect to yield, stage of development seemed to be of greater importance than the duration of waterlogging. The growth and yield of multiple-waterlogged sunflowers was less affected by the anthesis treatment than that in plants experiencing a single waterlogging, suggesting that some form of adaptation was induced. In contrast, no such response was seen in sorghum.     

Determinants of seed production in Geranium maculatum

Summary We experimentally examined factors limiting seed production in two populations of the perennial woodland herb Geranium maculatum in central Illinois, USA. To test the pollinator-limitation hypothesis, we compared the seed production of plants whose flowers were supplementarily pollinated with outcross pollen to that of control plants receiving natural pollination only. To test if fruit production by early flowers suppresses fruit and seed formation by late flowers, a third group of plants was prevented from producing seed from the first 50% of the flowers to open (stigmas were excised at flower opening). Finally, to test if seed maturation and flower initiation are correlated with photosynthetic capacity, we performed a defoliation experiment in which either the stem leaves within the inflorescence, the stem leaves below the inflorescence, or the rosette leaves were removed during late flowering. Plants that reccived supplemental pollination produced 1.5–1.6 times more seeds than control plants. We found no difference between hand-pollinated plants and controls in mortality, flowering frequency or number of flowers produced in the year following the experiment. In both control and hand-pollinated plants, the fruit set and total seed production of early flowers were more than twice as high as those of late flowers. In one of the two populations, plants whose early flowers were prevented from setting seed produced significantly more seeds from their late flowers than did control plants. Seed predation was low and did not differ between early and late flowers. Leaf removal did not affect seed number or size in the year of defoliation, nor did it reduce survival or flower production in the subsequent year. This suggests that the plants were able to compensate for a partial defoliation by using stored resources or by increasing photosynthetic rates in the remaining leaves. Taken together, the results demonstrate that both pollinator activity and resource levels influence patterns of seed production in G. maculatum. While seed production was pollinatorlimited in both populations, a seasonal decline in resource availability was apparently responsible for the low seed production by late flowers.     

A role for young leaves in vernalization of cauliflower: I. Analysis of leaf development during curd induction

Growth analysis techniques are used to test the hypothesis that chilling induces curd (flower) initiation in the cauliflower ( Brassica oleracea Botrytis L. cv. Perfection) through inhibiting leaf growth, thereby increasing the availability of growth factors to the stem apex and enabling differentiation of the curd. Effects of chilling on leaf growth and curd induction are compared in juvenile and mature, vegetative plants. Chilling at 5°C reduced dry matter accumulation in the total leaf complement by ca 60% in juvenile plants and 40% in mature plants, compared to control plants growth at 20°C. Juvenile plants showed slower rates of leaf initiation than mature plants. Leaf initiation was retarded by chilling in both plant types with the most marked effect seen in the juvenile plants. This was consistent with dry matter availability to the stem apex limiting differentiation more severely in juvenile plants than in mature plants. The rate of dry matter accumulation in existing leaves, however, was faster in juvenile plants than in mature plants at 20°C. Plants that were juvenile during chilling produced an average of 43 leaves below the curd whereas those that were mature produced 25.
Dry matter accumulation in younger leaves was more markedly inhibited by chilling than in older leaves. Chilling also reduced the rate at which enlarging leaves became positionally more remote from the stem apex. Possible roles for such leaves in regulating apical development are considered.     

The ecology and genetics of fitness in forest plants. IV. Quantitative genetics of fitness components in Impatiens pallida (Balsaminaceae)

Genetic and environmental variances were estimated for a number of characters in the annual plant Impatiens pallida by planting seed obtained through controlled crosses into their native field site or pots maintained in the greenhouse. Significant additive genetic variance was detected for three of 11 characters studied—germination date, cotyledon area, and date of first flower production. Significant dominance and/or maternal variance was found for seed weight, proportion of seeds germinating, cotyledon area, plant height, and number of leaves produced. Environmental variance was greater in the field compared with the greenhouse. Characters found to be under strong directional selection in a previous study showed no detectable additive genetic variance. While these populations exhibit conditions that in theory could contribute to the maintenance of genetic variation (limited pollen and seed dispersal distances and small-scale variation for edaphic characteristics influencing plant growth), levels of additive genetic variance for most characters were not significantly different from zero.     

Effects of Shading the First Leaf on Growth of Barley Plants: I. Long-term Experiments

Plants of barley were grown under controlled conditions andthe first or second leaves covered with tubular shades thusreducing the light intensity at the leaf surface to low levels.Expansion of the shaded leaves was not prevented, but appearanceof the next leaf but one and all subsequent leaves on the mainstemwas delayed by up to 3 days. Primordia of the first four leaveswere present in the dry grain. Shade treatment delayed slightlythe initiation of the eighth and subsequent leaves and transitionto the double ridge stage at the mainstem apex. Shading the first leaf caused a temporary reduction in the rateof dry-matter increase of plants, but after 14 days the ratewas similar to that of control plants. Smaller effects werefound when the second leaf was shaded. Dry-matter productionfollowed two logarithmic phases in the period prior to awn emergence,and rates for the whole plant and for plant parts were similarfor control and shaded plants. Thus, apart from the initialperturbation, shading had no effect on growth in terms of rateof dry-weight gain. Shade treatment did not affect weight per grain or numbers ofgrain per ear, but over-all yield of grain was significantlyreduced since shading delayed the appearance of tillers andalso reduced the number of tillers bearing grain. The effectof shade was especially marked on tillers originating on primarytillers. Similar qualitative effects on tiller development werefound in an experiment on wheat.     

Genetic divergence and biology of adaptation inCicer arietinum L.

Summary The role of 19 structural, developmental and biochemical traits in relation to specific adaptation was analysed in a set of 17 diverse lines with quantified adaptation, representing contemporary cultivars and land races of chickpea (Cicer arietinum L.), using multivariate analysis. Significant varietal variation was observed for most characters, particularly for the activity of the enzyme nitrate reductase (NR) and protein content in the plant. The distance analysis (D²-statistic) revealed that seed size and pod number and their associated attributes were important forces of divergence. The additional forces of divergence were NR activity at the flower initiation stage, yield components such as number of primary and secondary branches, and other features such as plant habit and duration of flowering. The principal component analysis revealed some similarities and also differences from the distance analysis. Leaf size, days to flower initiation, seed size and, to some extent, NR activity at flower initiation stage, were important in the first vector. Developmental traits such as chlorophyll depth, NR activity at the pod initiation and grain filling stages, and the percent protein content in the plant at flower initiation were important in the second vector. In general, the clustering pattern was not related to the geographical origin, seed colour, size of regression coefficient for yield, or deviation from linearity. The importance of the developmental and biochemical attributes in the divergence of cultivated chickpea, such as days to flower initiation, duration of flowering, NR activity and the rates of protein accumulation in developing seeds, and in adaptation, suggests the critical role of these attributes. NR activity at the flower initiation stage would appear to have a major role in the domestication of this crop and its intra-specific differentiation, as an increased seed size could not have been possible without better nutrient uptake and utilization.Part of dissertation submitted for Ph. D. Degree in genetics at the Post Graduate School, I.A.R.I., New Delhi in 1979, by R. G. Dani     

Opium poppy capsule growth and alkaloid production is constrained by shade during early floral development

Morphinan alkaloids accumulate in the capsules of Papaver somniferum L. (opium poppy) most likely as defence against herbivory. Thus, capsule size is an important component of alkaloid yield. Shade during early cell division-dominated growth of reproductive structures generally reduces final fruit size more than shade during later cell expansion-dominated growth. The current study aimed to determine whether this response is found in opium poppy and the subsequent impact on alkaloid yield, composition and seed production. First the timing of key reproductive developmental events was resolved relative to macromorphological traits. Plants were then shaded during either (a) floral initiation, (b) early floral development or (c) capsule expansion before being harvested at maturation. Shade during floral initiation dramatically reduced final capsule size, alkaloid yield and seed number, and increased the concentration of precursor compounds relative to morphine, despite plants later returning to full sun. Shade during later capsule growth enhanced capsule size and alkaloid yield but had little effect on alkaloid composition or seed number. Thus, early developmental processes, including morphine biosynthesis, appear to have a relatively greater demand for carbohydrates compared with later processes. Crop management practices and environmental factors that limit carbohydrate availability during early development are thus predicted to have significant negative impacts on alkaloid production and reproductive success.     

Effects of daylength and day/night temperatures on growth and seed yield of cowpea cv. K 2809 grown in controlled environments

The effects of climatic factors on the growth, reproductive development and seed yield of cowpea (cv. K 2809) were investigated in controlled environment cabinets. Plants were grown to maturity in eight environments comprising all combinations of two day lengths (11 h 40 min and 13 h 20 min), two day (27 and 33 ^oC) and two night (19 and 24 ^oC) temperatures. The plants were nodulated (Rhizobium strain CB 756) and received 197 ppm N throughout growth. Treatments changed the time to the appearance of first flowers by a maximum of 6 days but the later-flowering plants more than doubled their dry weight during this period, so that effects on plant form and, ultimately, seed production were considerable. Warm nights (24 ^oC) not only hastened the onset of flowering but also enhanced dry matter production during the pre-flowering period; they did not extend the total growing period. Warm days (33 ^oC) did not enhance dry matter production but shortened the duration of the growing period by an average of 21 days (20%). Variation in final seed yield was mainly due to differences in the number of pods borne on branches. Warm days markedly decreased the number of pods per plant (an overall average reduction of 49%) as did warm nights in conjunction with the long (13 h 20) daylength. The number of seeds per pod was effected only by daylength (8.3 and 7.6 seeds in the long and short daylengths, respectively). Mean seed weight was decreased by 19% in warm as compared to cool nights but was increased by 18% in warm as compared to cool days. These responses are compared with those obtained with soyabean cv. TK5 in a previous experiment and are shown, in general, to be similar.     

Pollination thoroughness and maternal yield regulation in wild radish,Raphanus raphanistrum (Brassicaceae)

Summary Under conditions where resources are limited, there are often negative correlations between components of maternal yield, or between fruit and flower production. Pollination, in turn, may vary among individuals and influence total maternal expenditure. We examined the impact of variation in pollination thoroughness upon yield components and overall plant growth in wild radish (R. raphanistrum) plants grown in the greenhouse. Plants received different pollination treatments in which 0% to 100% of all flowers produced were hand-pollinated. Fruit set was increased by hand-pollination, but rarely exceeded 30%, even when more than 50% of the flowers were pollinated. Plants receiving more thorough pollination or having greater proportion fruit set produced significantly smaller seeds. Seed number per fruit was not influenced by pollination treatment. Mean values of yield components and interactions between components often varied among plants from different maternal families. Increasing pollination thoroughness also resulted in dramatic decreases in flower production. If male fitness is related to flower number, there may be a tradeoff between maternal fecundity and successful pollen export operating at the whole-plant level in this species.     

Varying phosphorus supply and development,growth and seed yield in narrow-leafed lupin

Narrow-leafed lupin (Lupinus angustifolius L.) is usually grown in sandy, acidic and phosphorus (P) deficient soil with low yield and variable harvest index. This study aimed to examine the effects of varying P supply on lupin growth, seed yield and harvest index. Non-abscission plants (cv. Danja) were grown in Lancelin sand at seven rates of P supply (5, 10, 15, 20, 25, 30 or 40 mg kg^–1) in a naturally-lit glasshouse. The rate of leaf emergence, flowering time and flower number were decreased or delayed by low P supply (5, 10 or 15 mg kg^–1), with no differences at P rates higher than 20 mg kg^–1. High P supply (25, 30 or 40 mg kg^–1) increased plant seed yield and harvest index largely by increasing the number of pods and consequently yield on the lateral branches, but had less effect on the number of seeds per pod and seed size. Seed yield and seed P concentration continued to increase up to 40 mg P kg^–1but harvest index plateaued at 25 mg P kg^–1, indicating that low P supply decreased reproductive growth more than vegetative growth in narrow-leafed lupin.     

Effect of different constant soil moisture levels on foliage development in maize

The effects of constant soil moisture levels of 90, 60 and 40% of the maximum capillary capacity, applied beginning from the planting of the germinated kernels, on the dynamics of the foliage development of maize was studied in relation to ontogenical changes in leaf area of individual leaves. There were two maxima in the growth of total leaf area (characterized by leaf area duration and loaf area relative growth rate) unrelated to the soil mositure. The first maximum which was less marked, appeared in the phase of 5–8 leaves, the second and main increase being observed before flowering in the phase of 10–12 leaves. The effects of continued decrease in soil moisture were a correspondingly progressive reduction in leaf area and delayed development of the whole plants. Plants with 60% soil moisture attained the same leaf area as those with 90% but only after the end of the vegetative phase. The greatest differences in rate of development between watering treatments appeared at the time of greatest increase in leaf area of control plants. Continued decrease in soil moisture markedly affected the development of the leaves at different insertion levels (especially in elder leaves). Clear-cut prolongation of ontogenesis took place under dry conditions. In conditions of higher soil moisture growth was rather limited or stopped after reaching a certain maximum.     

Effects of Number of Photoperiodic Cycles on Induction and Development of Flowers and Fruits in Corchorus olitorius L. (var. oniyaya Epen.)

Plants of Corchorus olitorius, a short-day plant, were subjectedto varying numbers of short-day cycles before transfer to longdays. Treatments started after germination of the seeds at thetime of cotyledon release. Four short-day cycles (10 h naturaldaylight followed by 14 h darkness) were sufficient to induceflowering in all plants. The number of flowers and fruits producedon a plant increased as the number of short-day cycles was increasedfrom three to 30. Plants given three to seven short-day cyclesproduced flowers on the main stem only but when plants weremaintained in short days for longer periods, flowers were alsoproduced on the branches. The growth in d. wt of fruits wasgreatest when plants were maintained in short days throughout. Corchorus olitorius L., flowering, fruit development, photoperiodism