Post‐flowering night respiration and altered sink activity account for high night temperature‐induced grain yield and quality loss in rice (Oryza sativa L.)

High night temperature (HNT) is a major constraint to sustaining global rice production under future climate. Physiological and biochemical mechanisms were elucidated for HNT‐induced grain yield and quality loss in rice. Contrasting rice cultivars (N22, tolerant; Gharib, susceptible; IR64, high yielding with superior grain quality) were tested under control (23°C) and HNT (29°C) using unique field‐based tents from panicle initiation till physiological maturity. HNT affected 1000 grain weight, grain yield, grain chalk and amylose content in Gharib and IR64. HNT increased night respiration (Rn) accounted for higher carbon losses during post‐flowering phase. Gharib and IR64 recorded 16 and 9% yield reduction with a 63 and 35% increase in average post‐flowering Rn under HNT, respectively. HNT altered sugar accumulation in the rachis and spikelets across the cultivars with Gharib and IR64 recording higher sugar accumulation in the rachis. HNT reduced panicle starch content in Gharib (22%) and IR64 (11%) at physiological maturity, but not in the tolerant N22. At the enzymatic level, HNT reduced sink strength with lower cell wall invertase and sucrose synthase activity in Gharib and IR64, which affected starch accumulation in the developing grain, thereby reducing grain weight and quality. Interestingly, N22 recorded lower Rn‐mediated carbon losses and minimum impact on sink strength under HNT. Mechanistic responses identified will facilitate crop models to precisely estimate HNT‐induced damage under future warming scenarios.     

Carbon balance and source‐sink metabolic changes in winter wheat exposed to high night‐time temperature

Carbon loss under high night‐time temperature (HNT) leads to significant reduction in wheat yield. Growth chamber studies were carried out using six winter wheat genotypes, to unravel postheading HNT (23°C)–induced alterations in carbon balance, source‐sink metabolic changes, yield, and yield‐related traits compared with control (15°C) conditions. Four of the six tested genotypes recorded a significant increase in night respiration after 4 days of HNT exposure, with all the cultivars regulating carbon loss and demonstrating different degree of acclimation to extended HNT exposure. Metabolite profiling indicated carbohydrate metabolism in spikes and activation of the TriCarboxylic Acid (TCA) cycle in leaves as important pathways operating under HNT exposure. A significant increase in sugars, sugar‐alcohols, and phosphate in spikes of the tolerant genotype (Tascosa) indicated osmolytes and membrane protective mechanisms acting against HNT damage. Enhanced night respiration under HNT resulted in higher accumulation of TCA cycle intermediates like isocitrate and fumarate in leaves of the susceptible genotype (TX86A5606). Lower grain number due to lesser productive spikes and reduced grain weight due to shorter grain‐filling duration determined HNT‐induced yield loss in winter wheat. Traits and mechanisms identified will help catalyze the development of physiological and metabolic markers for breeding HNT‐tolerant wheat.     

Physiological factors limit fruit set of tomato (Lycopersicon esculentum Mill.) under chronic, mild heat stress

The effects of chronic, mild heat stress on fruit set, fruit production, release of pollen grains, photosynthesis, night respiration and anther dehiscence were examined in tomatoes (Lycopersicon esculentum Mill.) differing in high‐temperature sensitivity. Plants were grown under three temperature regimes: (1) 28/22 or 26/22 °C (optimal temperature); (2) 32/26 °C (high temperature); and (3) 32/26 °C day/night temperatures relieved at 28/22 °C for 10 d before anthesis, then returned to 32/26 °C (relieving treatment). FLA 7156 was the only cultivar with fruit set at 32/26 °C. All five cultivars, however, had fruit set under the relieving treatment (RT). The longer the relief, the higher the percentage of fruit set. Longer periods of relief also increased the number of pollen grains released, and linear regression analysis showed a significant relationship between the number of pollen grains released and the percentage of fruit set. Germination of pollen grains was also lowered in high‐temperature‐grown plants. The number of pollen grains produced, photosynthesis and night respiration did not limit fruit set under chronic, mild heat stress, however. This suggested that cultivar differences in pollen release and germination under heat stress are the most important factors determining their ability to set fruit.     

Leaf senescence and grain filling affected by post-anthesis high temperatures in two different wheat cultivars

High temperature is a major factor affecting grain yield and plant senescence in wheat growing regions of central and east China. In this study, two different wheat cultivars, Yangmai 9 with low-grain protein concentration and Xuzhou 26 with high-grain protein concentration, were exposed to different temperature regimes in growth chambers during grain filling. Four day/night temperature regimes of 34°C/22°C, 32°C/24°C, 26°C/14°C, and 24°C/16°C were established to obtain two daily temperatures of 28 and 20°C, and two diurnal day/night temperature differences of 12 and 8°C. Concentration of a lipid peroxidation product malondialdehyde (MDA), activities of the antioxidants superoxide dismutase (SOD) and catalase (CAT), chlorophyll concentration (SPAD) in flag leaves and kernel weight were determined. Results show that activities of SOD and CAT in leaves increased markedly on 14 days after anthesis (DAA) for the high-temperature treatment (34°C/22°C) and then declined. As a result, MDA concentration in leaves increased significantly under high temperature (34°C/22°C and 32°C/24°C). Compared with optimum temperature treatment, high temperature reduced the concentration of soluble protein and SPAD values in flag leaves. Grain-filling rate increased slightly initially, but decreased significantly during late grain filling under high temperature. As a result, final grain weight was reduced markedly under high temperature. Decreases in the activities of SOD and CAT and increases in MDA concentration in leaves were more pronounced with a 12°C of day/night temperature difference when under high temperatures. Kernel weight was higher under 12°C of day/night temperature difference under optimum temperatures (24°C/16°C and 26°C/14°C). The responses to high-temperature regimes appeared to differ between the two wheat cultivars with different grain protein concentrations. It is concluded that a larger diurnal temperature difference hastened the senescence of flag leaves under high-temperature conditions, but retarded senescence under optimum temperature treatments of 26°C/14°C and 24°C/16°C.     

Diurnal export and carbon economy in an expanding source leaf of cucumber at contrasting source and sink temperature

Effects of contrasting temperatures of an expanding leaf (source) and of remaining plant parts (sink) on diurnal export and distribution of carbon were studied in seedlings of Cucumis sativus L., cv. Farbio. The time course of the rate of export was calculated by measuring simultaneously the exchange of ¹⁴CO₂ and the amount of ¹⁴C in the source leaf by means of a Geiger-Müller detector using a steady-state labelling technique. In all treatments average export rate during the night (16 h) was maximally 50% of the average rate during the 8-h day. Temperature affected the diurnal course of export via the source leaf and the sink in different ways. At a source leaf temperature of 25 or 30°C export stopped 12 h after start of the night, whereas at 20°C export continued throughout the night. However, the total amount of carbon exported during a 24 h cycle, expressed as a proportion of the amount of carbon assimilated, was the same at source leaf temperatures of 20 or 30°C. Thus source leaf temperature did not affect the distribution of assimilates between source and sink, in contrast to sink temperature. After 24 h at a sink temperature of 30°C, 20% more ¹⁴C was exported to plant parts below the source leaf than with a sink temperature of 20°C, at the expense of carbon remaining in the source. During the day less starch and more structural dry matter was formed at a source leaf temperature of 30°C than at 20°C. After a complete day/night cycle, however, there was no difference between the treatments. Starch was the primary carbon source during the night, and the decline in the rate of export coincided with the depletion of starch. Thus the decline in the rate of export at a source leaf temperature of 25 or 30°C at 12 h after the start of the night was due to the depletion of starch at that time. Similarly, at 20°C export could continue until the end of the night as the starch degradation supplied assimilates during the whole night.     

Super-optimal temperatures are detrimental to peanut (Arachis hypogaea L.) reproductive processes and yield at both ambient and elevated carbon dioxide

Continuing increases in atmospheric carbon dioxide concentration (CO₂) will likely be accompanied by global warming. Our research objectives were (a) to determine the effects of season‐long exposure to daytime maximum/nighttime minimum temperatures of 32/22, 36/26, 40/30 and 44/34°C at ambient (350 μmol mol^?1) and elevated (700 μmol mol^?1) CO₂ on reproductive processes and yield of peanut, and (b) to evaluate whether the higher photosynthetic rates and vegetative growth at elevated CO₂ will negate the detrimental effects of high temperature on reproductive processes and yield. Doubling of CO₂ increased leaf photosynthesis and seed yield by 27% and 30%, respectively, averaged across all temperatures. There were no effects of elevated CO₂ on pollen viability, seed‐set, seed number per pod, seed size, harvest index or shelling percentage. At ambient CO₂, seed yield decreased progressively by 14%, 59% and 90% as temperature increased from 32/22 to 36/26, 40/30 and 44/34°C, respectively. Similar percentage decreases in seed yield occurred at temperatures above 32/22°C at elevated CO₂ despite greater photosynthesis and vegetative growth. Decreased seed yields at high temperature were a result of lower seed‐set due to poor pollen viability, and smaller seed size due to decreased seed growth rates and decreased shelling percentages. Seed harvest index decreased from 0.41 to 0.05 as temperature increased from 32/22 to 44/34°C under both ambient and elevated CO₂. We conclude that there are no beneficial interactions between elevated CO₂ and temperature, and that seed yield of peanut will decrease under future warmer climates, particularly in regions where present temperatures are near or above optimum.     

High night temperatures during grain number determination reduce wheat and barley grain yield: a field study

Warm nights are a widespread predicted feature of climate change. This study investigated the impact of high night temperatures during the critical period for grain yield determination in wheat and barley crops under field conditions, assessing the effects on development, growth and partitioning crop‐level processes driving grain number per unit area (GN). Experiments combined: (i) two contrasting radiation and temperature environments: late sowing in 2011 and early sowing in 2013, (ii) two well‐adapted crops with similar phenology: bread wheat and two‐row malting barley and (iii) two temperature regimes: ambient and high night temperatures. The night temperature increase (ca. 3.9 °C in both crops and growing seasons) was achieved using purpose‐built heating chambers placed on the crop at 19:000 hours and removed at 7:00 hours every day from the third detectable stem node to 10 days post‐flowering. Across growing seasons and crops, the average minimum temperature during the critical period ranged from 11.2 to 17.2 °C. Wheat and barley grain yield were similarly reduced under warm nights (ca. 7% °C^?1), due to GN reductions (ca. 6% °C^?1) linked to a lower number of spikes per m². An accelerated development under high night temperatures led to a shorter critical period duration, reducing solar radiation capture with negative consequences for biomass production, GN and therefore, grain yield. The information generated could be used as a starting point to design management and/or breeding strategies to improve crop adaptation facing climate change.     

Grain growth and endosperm cell size under high night temperatures in rice (Oryza sativa L.)

BACKGROUND AND AIMS: High night temperatures are more harmful to grain weight in rice than high day temperatures. Grain growth rate and growth duration were investigated to determine which was the cause of the decrease in final grain weight under high night temperatures. Endosperm cell number and cell sizes were also examined to determine which might cause the decrease in final grain weight. METHODS: Rice plants were grown outdoors in plastic pots and moved at heading time to three temperature-controlled glasshouses under high night temperature (HNT; 22/34 degrees C), high day temperature (HDT; 34/22 degrees C) and control conditions (CONT; 22/22 degrees C). Grains were sampled periodically, and the time-course of grain growth was divided into rate and duration by logistic regression analysis. Endosperm cell numbers and cell sizes were analysed by digitalized hand-tracing images of endosperm cross-sections. KEY RESULTS: The duration of grain growth was reduced by high temperature both day and night. However, the rate of grain growth was lower in HNT than in HDT. The number of cells in endosperm cross-sections in HNT was similar to that in HDT, and higher than that in CONT. The average cell area was smaller in HNT than in either CONT or HDT. The differences in average cell areas between HNT and HDT were greater at distances 60-80 % from the central point of endosperm towards the endosperm surface. CONCLUSIONS: The results show that HNT compared with HDT reduced the final grain weight by a reduction in grain growth rate in the early or middle stages of grain filling, and also reduced cell size midway between the central point and the surface of endosperm.     

Rice yield formation under high day and night temperatures—A prerequisite to ensure future food security

Increasing temperatures resulting from climate change dramatically impact rice crop production in Asia. Depending on the specific stage of rice development, heat stress reduces tiller/panicle number, decreases grain number per plant and lower grain weight, thus negatively impacting yield formation. Hence improving rice crop tolerance to heat stress in terms of sustaining yield stability under high day temperature (HDT), high night temperature (HNT), or combined high day and night temperature (HDNT) will bolster future food security. In this review article, we highlight the phenological alterations caused by heat and the underlying molecular-physiological and genetic mechanisms operating under different types of heat conditions (HDT, HNT, and HDNT) to understand heat tolerance. Based on our synthesis of HDT, HNT, and HDNT effects on rice yield components, we outline future breeding strategies to contribute to sustained food security under climate change.     

Photosynthetic rate and yield formation in different maize hybrids

The relationship between photosynthetic rate and yield formation processes of the newer and older maize hybrids were investigated. Leaf area at flowering (source) and kernel number (sink) of the newer hybrids were greater than the older ones although their light-saturated photosynthetic rate (P_sat) were not greater than the older ones before flowering. After flowering, P_sat and chlorophyll content of the newer hybrids declined more slowly than the older ones. They not only distributed almost all photosynthates produced after flowering to grain but also reallocated some reserved photosynthates produced before flowering to grain. The newer hybrids exhibited greater grain mass than the older ones mostly because they could optimally regulate the photosynthetic rate and yield formation processes to maximize grain mass.     

Effects of Asymmetric Heat on Grain Quality During the Panicle Initiation Stage in Contrasting Rice Genotypes

Heat stress during the panicle initiation stage hinders the formation of rice grains. It is speculated that heat exposure during the panicle initiation stage could influence grain quality in rice. To obtain preliminary knowledge on the effects of asymmetric heat on rice grain quality during the panicle initiation stage, four rice genotypes (Shanyou63, Liangyoupeijiu, Huanghuazhan, and Nagina22) were subjected to three heat treatments, i.e., high daytime temperature (HDT; 37.9 °C/24.5 °C), high nighttime temperature (HNT; 30.9 °C/30.5 °C), the combination of high daytime and nighttime temperature (HDNT; 38.5 °C/31.0 °C), and a control (CK, 31.5 °C/24.0 °C) in temperature-controlled greenhouses for 15 days during the panicle initiation stage. The milling and appearance qualities, which are crucial for commercial value, were studied. Heat treatments significantly reduced the amounts of brown rice, milled rice, and head rice and the grain length, grain width, chalky grain amount, and grain chalkiness in the rice genotypes Liangyoupeijiu, Nagina22, and Huanghuazhan during the panicle initiation stage, and the largest reductions in grain quality were frequently observed under HDNT treatment. The milling and appearance qualities in genotype Shanyou63 were negligibly affected by heat treatments and thus were regarded as tolerant to heat, and the rice genotypes Liangyoupeijiu, Huanghuazhan, and especially Nagina22 were susceptible to heat during the panicle initiation stage. We concluded that heat stress during panicle initiation impacted the milling and appearance qualities in rice, and differences existed among rice genotypes. The underlying mechanisms of the effects of heat on rice grain quality need further study.

     

High temperature effects on photosynthate partitioning and sugar metabolism during ear expansion in maize (Zea mays L.) genotypes

Short hot and dry spells before, or during, silking have an inordinately large effect on maize (Zea mays L.; corn) grain yield. New high yielding genotypes could be developed if the mechanism of yield loss were more fully understood and new assays developed. The aim here was to determine the effects of high temperature (35/27 °C) compared to cooler (25/18 °C) temperatures (day/night). Stress was applied for a 14 d-period during reproductive stages prior to silking. Effects on whole plant biomass, ear development, photosynthesis and carbohydrate metabolism were measured in both dent and sweet corn genotypes. Results showed that the whole plant biomass was increased by the high temperature. However, the response varied among plant parts; in leaves and culms weights were slightly increased or stable; cob weights decreased; and other ear parts of dent corn also decreased by high temperature. Photosynthetic activity was not affected by the treatments. The ¹³C export rate from an ear leaf was decreased by the high temperature treatment. The amount of ¹³C partitioning to the ears decreased more than to other plant parts by the high temperature. Within the ear decreases were greatest in the cob than the shank within an ear. Sugar concentrations in both hemicellulose and cellulose fractions of cobs in sweet corn were decreased by high temperature, and the hemicellulose fraction in the shank also decreased. In dent corn there was no reduction of sugar concentration except in the in cellulose fraction, suggesting that synthesis of cell-wall components is impaired by high temperatures. The high temperature treatment promoted the growth of vegetative plant parts but reduced ear expansion, particularly suppression of cob extensibility by impairing hemicellulose and cellulose synthesis through reduction of photosynthate supply. Therefore, plant biomass production was enhanced and grain yield reduced by the high temperature treatment due to effects on sink activity rather than source activity. Heat resistant ear development can be targeted for genetic improvement     

Responses of cow pea (Vigna unguiculata) varieties and their hybrids to variation in day and night temperature regimes

The effects of each of four contrasting combinations of two day (30, 26°C) and two night (24, 19°C) temperatures were studied on four cowpea varieties and their F₁ hybrids. Early vegetative development was enhanced by the higher day, higher night temperature and flowering occurred first in plants receiving this treatment. The hybrids showed marked heterosis for leaf area but not for time to flowering. The ten genotypes studied showed correlated responses to increases in night temperature and differed in the degree of their response. The extent of the response (or stability) of varieties was often determined by the character values in the environment with the highest yield and heterotic hybrids were the least stable. Grain yield was highest in the lower day temperature environments but there was variation between genotypes in whether this occurred at the lower or higher night temperature. The pattern of responses shown by hybrids agreed with those of their parents indicating that stability was an inherited character. The results confirmed and extended those in earlier studies and emphasised the importance of a greater understanding of the nature of responses to the environment in selecting new varieties.     

Heat-stress effects on reproduction and seed set in Linum usitatissimum L. (flax)

Heat stress can detrimentally affect the reproductive capacity of many plants. The effect of a 7 or 14 d heat stress on flowering, seed set, pollen viability and germinability of flax (Linum usistatissimum L.) was assessed under growth chamber conditions. An incremental (2 °C/h), cyclical (daytime high 40 °C and night‐time low 18 °C) heat stress was applied 12 d after the initiation of flowering. Although flower formation in flax was not affected by heat stress, boll formation and seed set were reduced with onset of the heat stress. On removal of heat stress the stressed plants showed a compensatory response, flowering and producing bolls at a greater rate than the control plants. Heat stress significantly prolonged flowering by 17 d. Boll weight and seed weight were reduced with heat stress and the number of malformed, sterile seed increased three‐fold after 14 d of heat stress. Pollen viability and appearance were negatively affected after 6 and 10 d of heat stress, respectively. Pollen germinability decreased by the sixth day of heat stress, with no pollen germinating by the tenth day. Effects of heat stress on pollen viability and germinability alone, which did not occur until after the sixth day of the stress, could not account for the decreased boll formation due to heat stress in flax. These observations suggest that a combined effect of heat stress on both pollen and ovules contributes to decreased boll formation and seed set in flax.     

Grain yield and kernel weight of two maize genotypes differing in nitrogen use efficiency at various levels of nitrogen and carbohydrate availability during flowering and grain filling

Grain yield per plant (GYP) and mean kernel weight (KW) of maize (Zea mays L.) are sensitive to changes in the environment during the lag phase of kernel growth (the time after pollination in which the potential kernel size is determined), and during the phase of linear kernel growth. The aim of this study was to assess genotypic differences in the response to environmental stresses associated with N and/or carbohydrate shortage at different phases during plant development. The rate and timing of N and carbohydrate supply were modified by application of fertilizer, shading, and varying the plant density at sowing, at silking or at 14 d after silking. The effects of these treatments on the photosynthetic capacity, grain yield and mean kernel weight were investigated in two hybrids differing in N use efficiency. The total above-ground biomass and grain yield per plant of the efficient hybrid responded little to altered environmental conditions such as suboptimal N supply, enhanced inter-plant competition, and shading for 14 d during flowering, when compared to the less efficient genotype. We conclude that grain yields in the efficient genotype are less sensitive not only to N stress, but also to carbohydrate shortage before grain filling. Shading of N deficient plants from 14 d after silking to maturity did not significantly reduce grain yield in the non-efficient genotype, indicating complete sink limitation of grain yield during grain filling. In the efficient genotype, in contrast, grain yield of N-deficient plants was significantly reduced by shading during grain filling. The rate of photosynthesis declined with decreasing foliar N content. No genotypic differences in photosynthesis were observed at high or low foliar N contents. However, at high plant density and low N supply, the leaf chlorophyll content after flowering in the efficient genotype was higher than that in the non-efficient genotype. Obviously, the higher source capacity of the efficient genotype was not due to higher photosynthetic N use efficiency but due to maintenance of high chlorophyll contents under stressful conditions. In the efficient genotype, the harvest index was not significantly affected by N fertilization, plant density, or shading before the grain filling period. In contrast, in the non-efficient genotype the harvest index was diminished by N deficiency and shading during flowering. We conclude that the high yielding ability of the efficient genotype under stressful conditions was associated with formation of a high sink capacity of the grains under conditions of low carbohydrate and N availability during flowering and with maintenance of high source strength during grain filling under conditions of high plant density and low N availability.     

Variation in responses to boron in rice

Background and aims

Boron (B) deficiency depresses grain set and grain yield of wheat and maize while having little effect on their vegetative growth. This paper describes effects of B deficiency in rice and how these vary with planting season and variety.

Methods

Three rice varieties (KDML105, CNT1, SPR1) were grown in sand culture without (B0) and with 10 μM (B10) B added to the nutrient solution, in the cool season of 2007/08 and 2008/09 and the hot season of 2011 in Chiang Mai, Thailand (18°47′N, 98°59′E). Boron responses were measured in growth and yield parameters, pollen viability and B concentration of the flag leaf and anthers at anthesis.

Results

Grain weight was strongly depressed by B deficiency ranging from 28 % in SPR1 to 79 % in CNT1, and the yield was much lower in the cool season than in the hot season plantings. The variation in grain weight was closely associated with grain set and number of spikelets but not with shoot dry weight or tillering. Grain set was closely related to pollen viability, and both were increased with increasing anther B concentration at >20 mg B kg^?1. In addition to its adverse effect on grain set, B deficiency also depressed grain filling and weight of individual grains in rice.

Conclusions

Boron deficiency depressed rice grain yield through adverse effects on reproductive growth, panicle and spikelet formation and grain filling, in addition to grain set as in wheat and maize.     

Effects of elevated temperature and carbon dioxide on seed-set and yield of kidney bean (Phaseolus vulgaris L.)

It is important to quantify and understand the consequences of elevated temperature and carbon dioxide (CO₂) on reproductive processes and yield to develop suitable agronomic or genetic management for future climates. The objectives of this research work were (a) to quantify the effects of elevated temperature and CO₂ on photosynthesis, pollen production, pollen viability, seed‐set, seed number, seeds per pod, seed size, seed yield and dry matter production of kidney bean and (b) to determine if deleterious effects of high temperature on reproductive processes and yield could be compensated by enhanced photosynthesis at elevated CO₂ levels. Red kidney bean cv. Montcalm was grown in controlled environments at day/night temperatures ranging from 28/18 to 40/30 °C under ambient (350 µmol mol^?1) or elevated (700 µmol mol^?1) CO₂ levels. There were strong negative relations between temperature over a range of 28/18–40/30 °C and seed‐set (slope, ? 6.5% °C^?1) and seed number per pod (? 0.34 °C^?1) under both ambient and elevated CO₂ levels. Exposure to temperature > 28/18 °C also reduced photosynthesis (? 0.3 and ? 0.9 µmol m^?2 s^?1 °C^?1), seed number (? 2.3 and ? 3.3 °C^?1) and seed yield (? 1.1 and ? 1.5 g plant^?1 °C^?1), at both the CO₂ levels (ambient and elevated, respectively). Reduced seed‐set and seed number at high temperatures was primarily owing to decreased pollen production and pollen viability. Elevated CO₂ did not affect seed size but temperature > 31/21 °C linearly reduced seed size by 0.07 g °C^?1. Elevated CO₂ increased photosynthesis and seed yield by approximately 50 and 24%, respectively. There was no beneficial interaction of CO₂ and temperature, and CO₂ enrichment did not offset the negative effects of high temperatures on reproductive processes and yield. In conclusion, even with beneficial effects of CO₂ enrichment, yield losses owing to high temperature (> 34/24 °C) are likely to occur, particularly if high temperatures coincide with sensitive stages of reproductive development.     

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.