Life history and photoperiodic responses in Chondrus giganteus forma flabellatus (Gigartinaceae,Rhodophyta) from Japan

The life history and photoperiodic responses of the carrageenophyte Chondrus giganteus forma flabellatus (Rhodophyta) from Japan were investigated in culture. This entity has a Polysiphonia-type life history and the formation of both tetrasporangia and cystocarps is controlled by daylength. A night-break of 1 h in the middle of a 16 h night was effective in inhibiting reproduction, confirming the classical photoperiodic nature of the response. Tetrasporangia formed at daylengths of 12 h or less, whereas cystocarps only developed at daylengths of 10 h or less. The formation of spermatangia required daylengths of 12 h or less, but production was not inhibited by a night-break of 1 h in the middle of a 16 h night.     

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.     

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.     

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.     

Time measurement and the control of flowering in plants

Many plants are adapted to flower at particular times of year, to ensure optimal pollination and seed maturation. In these plants flowering is controlled by environmental signals that reflect the changing seasons, particularly daylength and temperature. The response to daylength varies, so that plants isolated at higher latitudes tend to flower in response to long daylengths of spring and summer, while plants from lower latitudes avoid the extreme heat of summer by responding to short days. Such responses require a mechanism for measuring time, and the circadian clock that regulates daily rhythms in behaviour also acts as the timer in the measurement of daylength. Plants from high latitudes often also show an extreme response to temperature called vernalisation in which flowering is repressed until the plant is exposed to winter temperatures for an extended time. Genetic approaches in Arabidopsis have identified a number of genes that control vernalisation and daylength responses. These genes are described and models presented for how daylength might regulate flowering by controlling their expression by the circadian clock. BioEssays 22:38-47, 2000.     

Effects of Daylength on Floral Characteristics of Six White Clover (Trifolium repens) Varieties

The effect of daylength upon floral initiation, flower position,and associated features was measured in six white clover (Trifoliumrepens) varieties Most plants initiated flowers in the longest days (16 h). Thevarieties of more southerly origin and with large leaves hada greater spread of response to daylength. Increasing daylength increased the number of buds becoming inflorescencesin all varieties except Milkanova in which the reverse was true.Peduncle length and nectar production were both greatest inthe longest days. Trifolium repens, white clover, daylength, flower initiation, flower position     

Effects of daylength,irradiance and settlement density on the growth and reproduction of <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> gametophytes

The effects of daylength, irradiance and spore settlement density on the growth, maturation and sporophyte production of Undaria pinnatifida (Harvey) Suringar gametophytes were examined using a factorial experimental design in culture. The growth of Undaria gametophytes increased with increasing daylength (8, 12 and 16 h), but the maximum fertility occurred at a daylength of 12 h followed by 8 and 16 h. Gametophytes grew better at the 16 h daylength under the same mean daily irradiance (MDI) of 20 μmol photons m⁻² s⁻¹. However, the fertility was higher at the short daylength (8 h), indicating that the maturation of U. pinnatifida gametophytes is influenced by daylength rather than by the MDI. Vegetative growth and sporophyte production of gametophytes were better at 60 μmol photons m⁻² s⁻¹ than at 30 μmol photons m⁻² s⁻¹ under a 8:16 h LD (Light: Dark) cycle, and their growth and maturation were density-dependant in 16 and 12 h daylength, respectively. These results suggest that the U. pinnatifida gametophytes require a certain amount of light for the growth and reproduction, and intraspecific competition occurred under the optimal growth and maturation conditions. However, the sporophyte recruits per unit has been enhanced with increasing spore settlement density at 8 and 12 h daylengths indicating that high settlement density gives a benefit for maintaining population, even though the sporophyte production of each female plant is inhibited. In conclusion, the vegetative growth, reproduction and sporophyte production of U. pinnatifida gametophytes are retarded at a low irradiance above growth saturation and a high settlement density, and are determined by daylength.     

Reproductive response of a tropical mammal, the musk shrew (Suncus murinus), to photoperiod

Musk shrews (Suncus murinus) were maintained for 8 weeks in long (16 h light:8 h darkness) or short (8 h light:16 h darkness) daylengths. Males housed in short daylength had significantly lighter androgen-dependent sex accessory organs than did males kept in long daylengths. This same trend was noted in male sexual behaviour. However, the weights of the testes and epididymides and sperm numbers did not differ. Females housed in short daylengths had significantly lighter cervices and were less likely to demonstrate sex behaviour than animals kept in under long daylengths. Ovarian and uterine weights did not differ. These results suggest that the ability to respond to photoperiod can exist in tropical mammals, even if it is not used as a cue to time seasonal breeding.     

Morphology,temperature and photoperiodic responses in Audouinella botryocarpa (Harvey) Woelkerling (Acrochaetiaceae,Rhodophyta) from Ireland

Abstract

Audouinella botryocarpa is reported for the first time from the British Isles. On the west and south coasts of Ireland and in the Isle of Man it grows in the intertidal on wave-splashed limpet shells and rock surfaces. Populations examined monthly in Galway Bay formed monosporangia throughout the year and tetrasporangia from December to March and May but no gametangial plants were found. Plants isolated into culture from monospores formed monosporangia at daylengths of 8, 10, 12, 14 and 16 h at 6.5, 8, 10, 11 and 15°C; the same plants formed tetrasporangia at daylengths ≤10 h at 8, 10 and 11°C but not at 6.5 and 15°C. Spores isolated from these plants gave rise to further monosporangial plants that also formed tetrasporangia under inductive conditions. Night-breaks of 1 h in a 16 h night prevented tetrasporangial reproduction and a critical daylength of ～10 h was found at 10°C. British Isles Audouinella botryocarpa typically has 6 rounded or irregularly-shaped chloroplasts per cell, each of which has a single, centrally-placed pyrenoid. The chloroplasts grow together in mature cells, typically giving the appearance of a single plate-like chloroplast with 6 pyrenoids. The implications of these observations for the generic classification of acrochaetioid algae is discussed and it is concluded that chloroplast morphology, number and the presence or absence of pyrenoids may provide the basis for a future generic realignment of acrochaetioid algae.     

Influence of Ammonium on the Ability of Salicylic Acid to Induce Flowering in the Short-Day Plant Lemna paucicostata 6746

When the short-day plant Lemna paucicostata 6746 is grown inhalf-strength Hutner's medium, which contains 1.25 mM NH4NO3,salicylic acid does not induce flowering on daylengths of 16hr or longer. By contrast, in M (Hoagland-type medium), Pirson-Seidelor ammonium-free half-strength Hutner's media, none of whichcontain ammonium, salicylic acid is able to induce some floweringeven under continuous light. Neverthless, in each of these threemedia the effect of salicylic acid is strongly daylength dependentbecause there is a sharp drop in the flowering response to salicylicacid between the 12 and 16 hr daylengths, and the floweringresponse is nearly constant from the 16 hr daylength to continuouslight. Ammonium has the opposite effect and at 50 to 75 µMis able to overcome the salicylic acid effect completely andprevent any flowering on daylength of 16 hr or longer. (Received December 3, 1980; Accepted March 5, 1981)     

Photoperiodic control of larval development in the semivoltine cockroachPeriplaneta japonica (Blattidae: Dictyoptera)

Periplaneta japonica is semivoltine, entering early diapause in any (except the first) larval instar before the last, and late diapause in the last instar. Early diapause was induced under a short day of 13 h or less at 28°C, and under both short and long daylength (12–16 h) at 20°C. The shorter the daylength and the lower the temperature, the younger the instar was entering early diapause. Early diapause was terminated by a long day (16 h) or a high temperature (28°C), after which larvae grew faster in short days than in long days until the last instar, when they again entered diapause, always in short days and frequently in long days. This late diapause was terminated also by an increase in daylength and was always followed by adult emergence. In this case, 13 and 14 h daylengths after exposure to 12 h daylength were as effective as 16 h daylength. Ourdoor samples collected in late autumn, winter and early spring at Hirosaki (40.5°N) comprised two distinct size groups, corresponding with the early and late diapause instars. Based on these results, the seasonal development strategy and intriguing aspects of the photoperiodic response in this cockroach are discussed.     

Thermosensitive genetic dwarfs of apple

The role of environment on the dwarfing (short internode) phenomenon of apple (Malus domestisca Borkh.) was investi gated and defined in controlled environmental chambers. Orchard-grown very dwarf, dwarf and semi-dwarf trees obtained by natural sibcrossing of spur-type cv. Golden Delicious and cv. Delicious, as well as standard cv. Golden Delicious, were propagated via in vitro techniques. Growth was rapid and none of the 4 types exhibited dwarf-like characteristics when grown at constant 27°C with 12, 14 or 16 h daylengths. Standard and very dwarf plants grew at nearly the same rate at constant 30°C, whereas growth nearly ceased on both types at constant 35°C after 7 days. Dwarf and very dwarf plants responded differently from standard and semi-dwarf plants when grown under alternating (ramped) night/day temperatures (15 or 20°C night ramped up to a daytime maximum over 8 h of 23, 28, 33 or 38°C, held for 2 h and then ramped down over 5 h to the night temperature). As the night/maximum day temperature differentia) increased from 0 to 23° under the ramping environments, growth of dwarf plants decreased markedly as compared to standard plants. When the same night/maximum day temperature differential occurred, the effect on decreasing shoot length was greater at the higher (20°C) night temperature. Increasing maximum day temperatures under the ramped environment also reduced leaf area plant^?1 but did not markedly affect leaf number, resulting in short internodes. When a period of constant temperature was followed by ramped temperatures or vice versa, the sequence of constant vs ramped environments made little difference in the final growth of the 4 plant types. The data point to high temperature as the major factor for causing dwarfing of the sensitive plant types. Increasing the differential between night and maximum day temperature resulted in short internode. dwarf plants with small leaves similar to orchard-grown dwarf trees.     

Flowering of Stylosanthes guianensis in Relation to Juvenility and the Long-Short Day Requirement

Seedlings of Stylosanthes guianensis var. guianensis were grownin long (14 h) days in five temperature regimes for varyingperiods before transfer to short (11 h) days at 30 ?C/21 ?C.The juvenile phase before seedlings responded to inductive conditionswas c. 45–50 d, 50–60 d and 60–70 d for cv.Schofield, cv. Cook and C.P.I. 34906 respectively, which ispositively related to their critical photoperiod for flowering.Temperatures favourable for growth (e.g. 30 ?C/26 ?C) reducedthe juvenile phase in C.P.I. 34906 and in Cook, which did notflower in 11 h days unless previously exposed to more than 18long days. In a second experiment cv. Cook was confirmed as a long-shortday plant. Seedlings were grown for 50 d in a glasshouse withnatural daylength extended to 13, 14, 16 or 24 h before transferto 12 h photoperiods. Cook floral development was positivelyrelated to daylength provenance before transfer and plants incontinuous 12 h did not flower. Shortening daylength after 48 cycles of 12 h to 11.75 h didnot result in continued floral development in Cook plants butcv. Graham plants were initiated or transitional by 75 d. Key words: Stylosanthes guianensis, Photoperiod, Temperature, Flowering     

Abolition of seasonal testis changes in the Ile-de-France ram by short light cycles: relationship to luteinizing hormone and testosterone release

The purpose of this study was to examine the relationships between testis weight and the luteinizing hormone (LH) and testosterone releases in rams subjected to 14 successive 2-mo artificial light cycles in which daylength increased from 8 to 16 h in one month and decreased from 16 to 8 h the following month. Testis weights were measured fortnightly. Serial bleedings were performed during 6 to 8 h the last three 2-mo light cycles, when daylengths were 8, 12 (increasing), 16 and 12 h (decreasing) and plasma LH and testosterone were measured by radioimmunoassay. The overall mean testis weight, continuously high and equal to 314 g per testis, was not correlated with daylength. Mean LH levels and LH pulse height varied significantly according to daylength (P < 0.05 and P < 0.01, respectively) and were maximal on short days (8 h). Both mean and maximal testosterone levels presented variations according to daylength (P = 0.05). LH and testosterone pulsatility were quite high (equivalent to 7 to 10.3 pulses/day); however, mean plasma testosterone levels remained low. It is proposed that in rams under 2-mo light cycles, frequent but short stimulation of LH release by decreasing daylength phases allows maximal testis weight, while the interruption of LH stimulation by increasing daylengths prevents overstimulation of testosterone that can inhibit the LH release by negative feedback. Rams a under 2-mo light regimen presented a persistent high testis weight, indicating that seasonality had been broken down in the Ile-de-France male which is normally a marked seasonal breeder.     

Interactions of temperature and photoperiod in the control of flowering of latitudinal and altitudinal populations of wild strawberry (Fragaria vesca)

Floral induction and development requirements of a range of latitudinal and altitudinal Norwegian populations of the wild strawberry Fragaria vesca L. have been studied in controlled environments. Rooted runner plants were exposed to a range of photoperiods and temperatures for 5 weeks for floral induction and then transferred to long day (LD) at 20°C for flower development. A pronounced interaction of temperature and photoperiod was shown in the control of flowering. At 9°C, flowers were initiated in both short day (SD) and LD conditions, at 15 and 18°C in SD only, whereas no initiation took place at 21°C regardless of daylength conditions. The critical photoperiod for SD floral induction was about 16 h and 14 h at 15 and 18°C, respectively, the induction being incomplete at 18°C. The optimal condition for floral induction was SD at 15°C. A minimum of 4 weeks of exposure to such optimal conditions was required. Although the populations varied significantly in their flowering performance, no clinal relationship was present between latitude of origin and critical photoperiod. Flower development of SD-induced plants was only marginally advanced by LD conditions, while inflorescence elongation and runnering were strongly enhanced by LD at this stage. The main shift in these responses took place at photoperiods between 16 and 17 h. Unlike all other populations studied, a high-latitude population from 70°N ('Alta') had an obligatory vernalization requirement. Although flowering and fruiting in its native Subarctic environment and after overwintering in the field in south Norway, this population did not flower in the laboratory in the absence of vernalization, even with 10 or 15 weeks of exposure to SD at 9°C. Flowering performance in the field likewise indicated a vernalization requirement of this high-latitude population.     

Opposite effects of daylength and temperature on flowering and summer dormancy of Poa bulbosa

BACKGROUND AND AIMS: The timing of flowering and summer dormancy induction plays a central role in the adaptation of Mediterranean geophytes to changes in the length of the growth season along rainfall gradients. Our aim was to analyse the role of the variation in the responses of flowering and summer dormancy to vernalization, daylength and growth temperature for the adaptation of Poa bulbosa, a perennial geophytic grass, to increasing aridity. METHODS: Flowering and dormancy were studied under controlled daylengths [9 h short day (SD) vs. 16 h long day (LD)] and temperatures (16/10, 22/16 and 28/22 degrees C day/night) in four ecotypes originating in arid, semi-arid and mesic habitats (110, 276 and 810 mm rain year(-1), respectively) and differing in flowering capacity under natural conditions: arid-flowering, semi-arid-flowering, semi-arid-non-flowering and mesic-non-flowering. KEY RESULTS: Flowering and dormancy were affected in opposite ways by daylength and growth temperature. Flowering occurred almost exclusively under SD. In contrast, plants became dormant much earlier under LD than under SD. In both daylengths, high temperature and pre-chilling (6 weeks at 5 degrees C) enhanced dormancy imposition, but inhibited or postponed flowering, respectively. Induction of flowering and dormancy in the different ecotypes showed differential responsiveness to daylength and temperature. Arid and semi-arid ecotypes had a higher proportion of flowering plants and flowering tillers as well as more panicles per plant than mesic ecotypes. 'Flowering' ecotypes entered dormancy earlier than 'non-flowering' ecotypes, while the more arid the site of ecotype origin, the earlier the ecotype entered dormancy. CONCLUSIONS: Variation in the flowering capacity of ecotypes differing in drought tolerance was interpreted as the result of balanced opposite effects of daylength and temperature on the flowering and dormancy processes.     

FLOWERING OF DICHONDRA IN RESPONSE TO TEMPERATURE AND DAYLENGTH

Experiments using controlled environment facilities showed that flowering of Dichondra repens was promoted by chilling plants at 10 C or below. Optimum length of the chilling period was 5–6 weeks. Unchilled plants did not flower. The flowering stimulus resulting from chilling was destroyed by temperatures above 21 C. Rate of flowering was increased by short days during chilling, but short days could not substitute for the chilling requirement. Optimum daylength for flower initiation following chilling was approximately 14 hr and the optimum temperature was approximately 15 C. Flower buds developed in leaf axils of primary stems and laterals, but stem apices remained vegetative. When the chilling requirement was met flowering continued indefinitely as the plants grew.     

Temperature response and flowering of white clover (Trifolium repens) varieties in controlled environments and the field

Floral initiation of 10 white clover varieties growing in three controlled day/night temperature regimes, 22“/10°C, 20°/10°C, 17°/10°C, was recorded. Effects of artificial soil heating on floral initiation of the same plants subsequently transferred to the field were also examined. In the controlled environments only a slight increase in day temperature (2–5°C) was necessary to significantly increase flowering. Defoliation at the three temperatures had contrasting effects on subsequent flower production. Results from the soil heating experiment suggested that increased temperature might compensate for short daylengths, by bringing forward reproductive bud initiation by 1 month. Soil heating increased the total number of inflorescences produced.     

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.     

Climatic flowering requirements of bipolar sedges, Carex spp., and the feasibility of their trans-equatorial migration by mountain-hopping

A selection of northern and southern hemisphere populations of a range of bipolar Carex species was grown in controlled environments for comparison of their temperature and daylength requirements for flowering. All studied species have basically the same dual induction requirements for flowering but differ quantitatively in their respective critical temperatures and daylengths for primary and secondary floral induction. Usually, low temperature or short days (SD), over a wider range of temperatures, were required for initiation of inflorescence primordia, whereas long days (LD) advanced but were not obligatory for heading and flower development. It is concluded that no change or adaptation in the flowering requirements would have been necessary for these plants to migrate across the tropics and their penetration into the southern hemisphere by mountain-hopping. The flowering requirements of these and a range of other bipolar plant species are compatible with the hypothesis that absence of an obligatory long day floral induction requirement has facilitated or been a prerequisite for trans-equatorial migration of plants by mountain-hopping and the establishment of bipolar species distributions.