An investigation of the frost hardiness of grapevine (Vitis vinifera) during bud break

The aim of this investigation was to assess ice nucleation and frost resistance of two varieties of grapevine (Siegrebbe and Madeleine Angevine) during bud burst under radiative freezing conditions analogous to those during Spring in the UK. During bud burst, grapevines were almost entirely resistant to freezing during frosts of less than -3°C by virtue of their ability to supercool. The risk of frost damage increased significantly as bud development progressed, and once buds had passed growth stage DS3 they became more sensitive to freezing and freezing damage was more extensive. The two varieties did not differ in frost resistance but, because of its earlier developing habit, variety Siegrebbe could be more prone to frost damage in the field. Buds were more prone to damage after freezing once bud burst had commenced and the damage could not be reversed by acclimating plants for periods of 7 to 21 days at 4°C in an 8 h photoperiod. Such acclimation appeared to predispose frozen buds to more extensive damage.     

Photosynthesis in frost-hardened and frost-stressed leaves of Hedera helix L.

Abstract The purpose of this study was to determine the respective extents to which winter reduction of photosynthetic capacity in ivy (Hedera helix L.) is caused by direct frost injury to the photosynthetic apparatus and by preceding protoplasmic changes connected with the acquisition of frost tolerance. Potted juvenile ivy plants were placed in the open under natural weather conditions whilst others were hardened under controlled conditions and subjected to the desired frost stress. Low non-freezing temperatures induced frost tolerance in ivy leaves down to about – 12°C (50% injury = TL₅₀) without impairing net photosynthetic rate as measured under standard conditions (20°C, light saturation, natural CO₂ level; = Standard-F_n. Only if the leaves froze (below ? 3°C to ?4°C) was a reversible inhibition of Standard-F_n observed. As long as the temperatures did not fall below approximately ?8°C the inhibition was small and Standard-F_n reached about 80–90% of the control. In this case the stomatal opening narrowed, giving a poorer supply of CO₂ to the mesophyll cells. Maximal frost tolerance (TL_5O?20°C to ?24°C) developed only with severe frosts below about ? 10°C. After such frosts, Standard-F_n was reduced to less than 20% of the control. The dependence of the rate of net photosynthesis on the internal CO₂ concentration showed a lower initial slope, thus indicating disturbances of chloroplast functions. However, neither in outdoor plants nor in those artificially frosted at – 20°C could there be found an appreciable inhibition of the electron transport capacity from H₂O to dichlorophenol indophenol or of ribulose bisphosphate carboxylase. If intact, severely frosted ivy plants were then held at higher temperatures (20/15°C), Standard-F_n recovered completely in approximately 10 d. Furthermore, following a frost period with temperatures down to ?12°C, mild weather caused a distinct improvement in Standard-F_n in outdoor plants, and there was no loss of maximum frost tolerance. Thus it can be concluded that the inhibition of Standard-F_n after severe frosts is not due to the development of maximal frost tolerance, but rather may be attributed to frost damage to the photosynthetic apparatus.     

The effect of LAB 173711 and ethephon on time of flowering and cold hardiness of peach flower buds

Peach flowers are often killed during bloom by spring frosts. LAB 173711, a compound with abscisic (ABA)-like activity, and ethephon delayed flowering in peach trees. In greenhouse experiments, LAB 173711, at concentrations of 10^?3–10^?2 M, was most effective in delaying bloom when applied after a 5°C cold storage period, rather than before the dormancy breaking treatment. In contrast, ethephon delayed bloom most effectively when applied before 5°C cold storage; ethephon caused flower bud abscission when treatments were made after the chilling requirement had been satisfied. In field experiments, ethephon delayed flowering by 6–7 days, which reduced bud injury after a spring frost during bloom. No flower bud injury was found on ethephon-treated trees after temperatures of ?4.3°C; whereas without ethephon 25% of the flower buds were frost damaged. LAB 173711 delayed the time to 50% bloom by 2–3 days. However, this was not long enough to avoid low-temperature injury to the flower buds.     

A chamber for the simulation of radiation freezing of plants

Frost injury to plants can occur following episodic radiation frosts. In the UK this is particularly important to spring sown crops such as potatoes. Most laboratory based frost studies simulate freezing using either conductive or convective freezing chambers. Such frost tests do not simulate overnight freezing events adequately. A freezing chamber based on radiative cooling is described which mimics overnight radiative freezing. The chamber is rectangular in design (1 m × lm × 2 m high) with a radiative cooling plate at the top of the chamber cooled to -40°C to -45°C using HFC coolants, which acts as a cold black body. The sides of the chamber are also cooled to variable temperatures down to -5°C in order to prevent the chamber walls radiating to the plant material during testing. Using thermocouples to measure air temperature and plant temperature the chamber has been characterised to simulate the radiative cooling conditions found in the UK during autumn and spring. Exotherm detection upon plant freezing is simplified by virtue of the reduction in temperature fluctuation normally experienced at the plant surface during natural freezing. Radiation frosts and subsequent frost damage to potatoes have been recorded in the temperature range -4°C to –5°C. The equipment is recommended for studies of frost damage to plants normally caused by episodic radiation frost events.     

Effect of thermal shock in the development of androgenic plants of <Emphasis Type="Italic">Anemone coronaria</Emphasis> L.: influence of genotype and flower parameters

Anemone is an allogamous species marketed as a cut flower. Pure lines formation in this species is expensive and inefficient because progeny suffers from inbreeding depression. Pure lines obtained from plant anthers are a widely used material to produce new hybrids in short time. The aim of this study is the optimization of production of A. coronaria androgenic plants. Flower buds of eight cultivars were measured, disinfected for 20′ in 1% sodium hypochlorite and rinsed with distilled sterile water. Development stage of microspores and their size were evaluated. Anthers were distributed in Petri plates containing a double layer constituted of Nitsch and Nitsch agarized substrate enriched with activated charcoal (1%) and with or without sucrose (3%) and the same substrate in liquid phase without activated charcoal. Anthers were shocked with a preculture at 5 or 33 °C for 5 days and then incubated in the dark at 23 °C. After 21 weeks, the number of regenerants was counted. Plants were transferred into the flask with modified MS substrate and placed at 18 °C. After 3 months of in vitro culture, the survival rate of plants was evaluated. Among the varieties, significant differences in anther number, microspore size, and presence of abnormal microspores were observed. Heat shock pretreatment at 33 °C increases the number and survival of embryos and plants, but the varieties respond to the thermal shock in different ways. The influence of genotype and the maturation phase of flower buds, anthers and microspores are discussed related to the androgenic plant development.     

The effect of LAB 173711 and ethephon on time of flowering and cold hardiness of peach flower buds

Peach flowers are often killed during bloom by spring frosts. LAB 173711, a compound with abscisic (ABA)-like activity, and ethephon delayed flowering in peach trees. In greenhouse experiments, LAB 173711, at concentrations of 10^–3–10^–2 M, was most effective in delaying bloom when applied after a 5°C cold storage period, rather than before the dormancy breaking treatment. In contrast, ethephon delayed bloom most effectively when applied before 5°C cold storage; ethephon caused flower bud abscission when treatments were made after the chilling requirement had been satisfied. In field experiments, ethephon delayed flowering by 6–7 days, which reduced bud injury after a spring frost during bloom. No flower bud injury was found on ethephon-treated trees after temperatures of –4.3°C; whereas without ethephon 25% of the flower buds were frost damaged. LAB 173711 delayed the time to 50% bloom by 2–3 days. However, this was not long enough to avoid low-temperature injury to the flower buds.     

How endangered is sexual reproduction of high-mountain plants by summer frosts? Frost resistance, frequency of frost events and risk assessment

In temperate-zone mountains, summer frosts usually occur during unpredictable cold spells with snow-falls. Earlier studies have shown that vegetative aboveground organs of most high-mountain plants tolerate extracellular ice in the active state. However, little is known about the impact of frost on reproductive development and reproductive success. In common plant species from the European Alps (Cerastium uniflorum, Loiseleuria procumbens, Ranunculus glacialis, Rhododendron ferrugineum, Saxifraga bryoides, S. moschata, S. caesia), differing in growth form, altitudinal distribution and phenology, frost resistance of reproductive and vegetative shoots was assessed in different reproductive stages. Intact plants were exposed to simulated night frosts between ?2 and ?14 °C in temperature-controlled freezers. Nucleation temperatures, freezing damage and subsequent reproductive success (fruit and seed set, seed germination) were determined. During all reproductive stages, reproductive shoots were significantly less frost resistant than vegetative shoots (mean difference for LT₅₀ ?4.2 ± 2.7 K). In most species, reproductive shoots were ice tolerant before bolting and during fruiting (mean LT₅₀ ?7 and ?5.7 °C), but were ice sensitive during bolting and anthesis (mean LT₅₀ around ?4 °C). Only R. glacialis remained ice tolerant during all reproductive stages. Frost injury in reproductive shoots usually led to full fruit loss. Reproductive success of frost-treated but undamaged shoots did not differ significantly from control values. Assessing the frost damage risk on the basis of summer frost frequency and frost resistance shows that, in the alpine zone, low-statured species are rarely endangered as long as they are protected by snow. The situation is different in the subnival and nival zone, where frost-sensitive reproductive shoots may become frost damaged even when covered by snow. Unprotected individuals are at high risk of suffering from frost damage, particularly at higher elevations. It appears that ice tolerance in reproductive structures is an advantage but not an absolute precondition for colonizing high altitudes with frequent frost events.     

Influence of stock plant pretreatment on gynogenic embryo induction from flower buds of onion

The gynogenic response of a range of onion genotypes to flower bud culture was compared using a two-step culture system. Embryogenic cultures and plantlets were produced from unpollinated ovules in whole flower bud explants 6 to 19 weeks after culture initiation. Preconditioning stock plants significantly influenced gynogenic embryogenesis. A ten-fold increase in embryogenesis was obtained when flower buds were cultured from stock plants maintained at 15 °C compared to 10 °C or the ambient temperature conditions of a glasshouse (maximum-minimum of 25–12.7 °C). A total of 49 embryos was obtained from 2660 cultured flower buds and 45% of plantlets were successfully acclimatised to glasshouse conditions. The majority of acclimatised plantlets were haploid (68%) but spontaneous double haploid plants (23%) were obtained from three genotypes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Critically reduced frost resistance of Picea abies during sprouting could be linked to cytological changes

Frost resistance of sprouting Picea abies shoots is insufficient for survival of naturally occurring late frosts. The cellular changes during sprouting appeared to be responsible for frost damage as frost events that damaged sprouting shoots did not damage older needles and stems. Whilst resting buds showed initial frost damage at ?15.0°C, 20 days later, current year’s growth was damaged at ?5.6°C. The decrease in frost resistance in sprouting shoots of P. abies was accompanied by a significant reduction of the cellular solute concentration, indicated by much less negative Ψ_oSAT values (increase from ?2.8 to ?1.2 MPa). ψ_oSAT decreased again after the final cell volume was reached and cell wall thickening began. After bud break, ice nucleation temperature increased from ?4.7°C to ?1.5°C. This increase was probably caused by the loss of bud scales, the onset of expansion growth of the central cylinder and the development of vascular tissue permitting the spread of ice from the stem into the growing needles. The onset of mesophyll cell wall thickening coincided with the lowest frost resistances. Cell wall thickening caused an increase in the modulus of elasticity, ε, indicating a decrease in tissue elasticity and after that frost resistance increased again. Metabolic and cytological changes that evidently leave little leeway for frost hardening are responsible for the low frost resistance in current year’s growth of P. abies. This low frost resistance will be significant in the future as the risk of frost damage due to earlier bud break is anticipated to even further increase.     

Flower bud differentiation in Salix pentandra as affected by photoperiod, temperature and growth regulators

Flower bud initiation in seedlings and vegetatively propagated plants of Salix pentandra from different locations has been studied under controlled conditions. In mature plants flower bud formation was induced by 2-chloroethyltrimethylammoniumchloride (CCC) and by short day treatment. The effect of CCC was antagonized by GA₃. The critical photoperiod for flower bud formation was about 18 h for a southern clone (from 49°48'N), but cuttings of a northern ecotype (from 69°39'N) formed flower buds even at 24 h photoperiod. Generally, flower bud formation occurred simultaneously with apical growth cessation. However, apical growth cessation was not a prerequisite for floral initiation and flower buds were also found in elongating plants. Seedlings up to 60 days old did not form flower buds in growth chamber studies. The length of the juvenile phase has not been studied in detail, but cuttings taken from seedlings approximately 20 cm high and 60 days old were able to develop flower buds when treated with CCC. A gradual transition from the juvenile to the mature phase was obtained by repeated pruning of seedlings grown at 18°C and 24 h photoperiod.     

The ecological and evolutionary significance of frost in the context of climate change

The effects that below-freezing temperature (frost) can have at times of year when it is unusual are an interesting ecological phenomenon that has received little attention. The physiological consequence of formation of ice crystals in plant tissue is often death of the plants, or at least of sensitive parts that can include flower buds, ovaries, and leaves. The loss of potential for sexual reproduction can have long-lasting effects on the demography of annuals and long-lived perennials, because the short-term negative effects of frosts can result in longer-term benefits through lowered populations of seed predators. The loss of host plants can have dramatic consequences for herbivores, even causing local extinctions, and the loss of just flowers can also affect populations of seed predators and their parasitoids. Frosts can cause local extinctions and influence the geographical distribution of some species. The potential for global climate change to influence the frequency and distribution of frost events is uncertain, but it seems likely that they may become more frequent in some areas and less frequent in others.     

Freezing Avoidance Mechanisms by Supercooling in Some Rhododendron flower Buds with Reference to Water Relations

Excised florets of some hardy Rhododendron species did not toleratefreezing at –5°C when ice-inoculated due to intracellularfreezing. Florets in intact December buds, however, could besupercooled to about –30°C. When flower buds of R.japonicum were slowly cooled with daily decrements of 5°Cto temperatures ranging from 0 to –20°C, the exothermtemperatures of the florets drastically decreased. This wasaccompanied by a decrease in water content of florets and peduncleand an increase in that of scales. The water in florets andthe peduncle is thought to migrate to scales and other tissuesduring the early stages of freezing; the dehydrated floret hasa lower freezing point which enhances its supercooling abilityand the dehydrated peduncle helps to maintain the supercooledstate of the florets. This hypothesis would explain the dependenceon the cooling rate of supercooling in Rhododendron flower buds.Water migration within flower buds was observed in other hardyRhododendron species with some variation in ice formation siteand the quantity of migrated water. The exotherm temperatureof excised florets was inversely proportional to their watercontent. Dehydration of flower buds by wind at 0°C alsoenhanced their supercooling ability. Mechanisms of freezingavoidance by supercooling in Rhododendron flower buds and therelationship of supercooling to freezing tolerance are discussed. ¹ Contribution No. 2254 from the Institute of Low TemperatureScience ² This is a revised form of the master's thesis of the seniorauthor (M.I.) which is cited in the present and previous papers(Sakai 1979a, b, etc.). (Received August 11, 1980; Accepted June 1, 1981)     

Photosynthesis in cold acclimated leaves of plants with various degrees of freezing tolerance

The objective of this study was to compare the photosynthetic changes during cold acclimation in various plant types able to acquire different degrees of freezing tolerance. Four herbaceous and six woody plants were hardened under natural or artificial conditions and – after determination of their frost resistance (LT₅₀) – the net photosynthetic rate at an ambient CO₂ of 33 Pa (P_n33), the dependencies of P_n to light and to CO₂ and the room temperature chlorophyll a fluorescence were recorded under optimal conditions. Herbaceous plants acquired freezing tolerances to temperatures between ?10 and ?15°C when hardened at temperatures around 0°C. Most leaves fully developed prior to frost hardening exhibited typical symptoms of senescence after frost hardening. In non-senescing leaves P_n33 was reduced by 15 to 50% mainly due to a reduced stomatal conductance. After hardening at temperatures around ?10°C Brassica survived down to ?24°C, but P_n33 was almost abolished as a result of disturbances in the chloroplasts. After transferring the plants to 20/15°C P_n33 recovered completely within a few days. Woody plants hardened at temperatures around 0°C tolerated – 15 to ?36°C: P_n33 was reduced by 25 to 60% and hardly recovered at 20/15°C. Hardening at ?10°C induced a tolerance of ?32 to n33 was almost totally blocked, but at 20/15°C it returned to the values of the plants hardened at 0°C within a few days. In woody plants disturbances were invariably localized in the chloroplasts. Thus, conifers, and especially Pinus cembra, can survive much lower temperatures than herbaceous plants and, at the same level of freezing tolerance, exhibit appreciably less restriction in relative P_n33.     

Frost in a future climate: modelling interactive effects of warmer temperatures and rising atmospheric [CO2] on the incidence and severity of frost damage in a temperate evergreen (Eucalyptus pauciflora)

Although plants are more susceptible to frost damage under elevated atmospheric [CO₂], the importance of frost damage under future, warmer climate scenarios is unknown. Accordingly, we used a model to examine the incidence and severity of frost damage to snow gum (Eucalyptus pauciflora) in a sub‐alpine region of Australia for current and future conditions using the A2 IPCC elevated CO₂ and climate change scenario. An existing model for predicting frost effects on E. pauciflora seedlings was adapted to include effects of elevated [CO₂] on acclimation to freezing temperatures, calibrated with field data, and applied to a study region in Victoria using climate scenario data from CSIRO's Global Climate Model C‐CAM for current (1975–2004) and future (2035–2064) 30 years climate sequences. Temperatures below 0 °C were predicted to occur less frequently while the coldest temperatures (i.e. those below ?8 °C) were almost as common in the future as in the current climate. Both elevated [CO₂] and climate warming affected the timing and rates of acclimation and de‐acclimation of snow gum to freezing temperatures, potentially reducing the length of time that plants are fully frost tolerant and increasing the length of the growing season. Despite fewer days when temperatures fall below 0 °C in the future, with consequently fewer damaging frosts with lower average levels of impact, individual weather sequences resulting in widespread plant mortality may still occur. Furthermore, delayed acclimation due to either warming or rising [CO₂] combined with an early severe frost could lead to more frost damage and higher mortality than would occur in current conditions. Effects of elevated [CO₂] on frost damage were greater in autumn, while warming had more effect in spring. Thus, frost damage will continue to be a management issue for plantation and forest management in regions where frosts persist.     

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.     

Direct and indirect effects of episodic frost on plant growth and reproduction in subalpine wildflowers

Frost is an important episodic event that damages plant tissues through the formation of ice crystals at or below freezing temperatures. In montane regions, where climate change is expected to cause earlier snow melt but may not change the last frost‐free day of the year, plants that bud earlier might be directly impacted by frost through damage to flower buds and reproductive structures. However, the indirect effects of frost mediated through changes in plant–pollinator interactions have rarely been explored. We examined the direct and pollinator‐mediated indirect effects of frost on three wildflower species in southwestern Colorado, USA, Delphinium barbeyi (Ranunculaceae), Erigeron speciosus (Asteraceae), and Polemonium foliosissimum (Polemoniaceae), by simulating moderate (?1 to ?5°C) frost events in early spring in plants in situ. Subsequently, we measured plant growth, and upon flowering measured flower morphology and phenology. Throughout the flowering season, we monitored pollinator visitation and collected seeds to measure plant reproduction. We found that frost had species‐specific direct and indirect effects. Frost had direct effects on two of the three species. Frost significantly reduced flower size, total flowers produced, and seed production of Erigeron. Furthermore, frost reduced aboveground plant survival and seed production for Polemonium. However, we found no direct effects of frost on Delphinium. When we considered the indirect impacts of frost mediated through changes in pollinator visitation, one species, Erigeron, incurred indirect, negative effects of frost on plant reproduction through changes in floral traits and pollinator visitation, along with direct effects. Overall, we found that flowering plants exhibited species‐specific direct and pollinator‐mediated indirect responses to frost, thus suggesting that frost may play an important role in affecting plant communities under climate change.     

Deep supercooling enabled by surface impregnation with lipophilic substances explains the survival of overwintering buds at extreme freezing

The frost survival mechanism of vegetative buds of angiosperms was suggested to be extracellular freezing causing dehydration, elevated osmotic potential to prevent freezing. However, extreme dehydration would be needed to avoid freezing at the temperatures down to ?45°C encountered by many trees. Buds of Alnus alnobetula, in common with other frost hardy angiosperms, excrete a lipophilic substance, whose functional role remains unclear. Freezing of buds was studied by infrared thermography, psychrometry, and cryomicroscopy. Buds of A. alnobetula did not survive by extracellular ice tolerance but by deep supercooling, down to ?45°C. An internal ice barrier prevented ice penetration from the frozen stem into the bud. Cryomicroscopy revealed a new freezing mechanism. Until now, supercooled buds lost water towards ice masses that form in the subtending stem and/or bud scales. In A. alnobetula, ice forms harmlessly inside the bud between the supercooled leaves. This would immediately trigger intracellular freezing and kill the supercooled bud in other species. In A. alnobetula, lipophilic substances (triterpenoids and flavonoid aglycones) impregnate the surface of bud leaves. These prevent extrinsic ice nucleation so allowing supercooling. This suggests a means to protect forestry and agricultural crops from extrinsic ice nucleation allowing transient supercooling during night frosts.     

Effect of frost nights and day and night temperature during dormancy induction on frost hardiness, tolerance to cold storage and bud burst in seedlings of Norway spruce

For trees, the ability to obtain and maintain sufficient levels of frost hardiness in late autumn, winter and spring is crucial. We report that temperatures during dormancy induction influence bud set, frost hardiness, tolerance to cold storage, timing of bud burst and spring frost hardiness in seedlings of Norway spruce (Picea abies (L.) Karst.). Bud set occurred later in 12°C than in 21°C, and later in cool nights (7°C) than in constant temperature. One weekly frost night (−2.5°C) improved frost hardiness. Cool nights reduced frost hardiness early, but improved hardiness later during cold acclimation. Buds and stems were slightly hardier in 21°C than in 12°C, while needles were clearly hardier in 12°C. Cold daytime temperature, cool nights and one weekly frost night improved cold storability (0.7°C). Seedlings receiving high daytime temperatures burst buds later, and were less injured by light frost some days after bud burst.     

Alleviation of frost damage to pear flowers by application of gibberellin

Adverse effects of gibberellin applications on pear trees after frost such as small fruit size, abnormal fruit shape and poor return bloom are often attributed both to the sole use of GA₃ and its overdose. It is unclear whether protection against spring frosts before flower opening is more efficient when GA is applied directly after frost, i.e. before flower opening, or at full bloom or both. In April 2003, early spring frosts at Klein-Altendorf near Bonn, Germany damaged ca. 88% flowers of the early flowering cv. Alexander Lucas, 64% of cv. Conference and ca. 25% of flowers of the later flowering cv. Comice pears. Hence, the objective of the present work was to investigate the optimum timing of the application of low doses of the combined GA₃ and GA_{4 + 7} to improve parthenocarpic fruit set in pears, while maximising fruit quality and size for trees affected by a severe spring frost before full bloom. Return bloom was also considered and quantified. Frost-affected pear trees were treated with gibberellin GA₃+GA_{4 + 7}, either immediately after the frost, at the white bud stage, or at full bloom or both to improve parthenocarpic set. Early flowering cv. Alexander Lucas pear was most affected by the early spring frost, but lost only 25% of fruitlets at June drop, irrespective of GA treatment. June drop was, however, severe in the two other cultivars least affected by frost, i.e. by 33% in cv. Conference and 55% in cv. Comice. Both initial and final fruit set were significantly increased by a combined application of GA₃+GA_{4 + 7} at full bloom, without affecting return bloom, but June drop was also enhanced by GA application. The largest positive effect of GA application on fruit yield, an additional 2 kg of fruit per tree equivalent to €1200/ha, was apparent with the cv. Alexander Lucas, i.e. the cultivar most affected by frost. There was no loss in fruit quality viz fruit size after any of the GA applications with any of the pear cultivars examined and no increase in abnormally-shaped, elongated fruit.     

Environmental Factors Controlling Flower Opening and Closing in aPortulacaHybrid

To examine flower opening and closing of aPortulacahybrid, flowerbuds were placed in darkness for 12 h (2030–0830 h) at20 °C and then exposed to various light-temperature conditions.Flower buds exposed to light at 25, 30 or 35 °C opened within1 h, and wilted 10–14 h later. Flower buds exposed tolight at 20 °C started to open after 4 h but opened slowlyand not completely. Flower buds subjected to 25, 30 or 35 °Cin darkness also opened rapidly, but did not reach full opening.Flowers opened at 30 °C in light, and partially closed andopened repeatedly in response to cycles of a 2-h exposure to20 °C and a 2-h exposure to 30 °C at any time between1000 to 1600 h. Similar phenomena were observed when the flowersopened at 30 °C in light and then were subjected to darknessand light alternately at 30 °C, although the effect of lightwas less obvious than that of alternating temperature. Floweropening and closing were not affected by relative humidity.These results indicate that a rise in temperature is requiredfor rapid flower opening in the buds kept at 20 °C, andthat light intensifies the effect of high temperature. Exposureto light without a temperature change delayed and slowed floweropening which was never complete. The involvement of an endogenousrhythm in flower opening byPortulacais indicated. Portulacahybrid, flower opening, flower closing, temperature shift, endogenous rhythm.