The formation and distribution of ice within dormant and deacclimated peach flower buds

Abstract A freeze-fixation technique was used to examine the distribution of ice crystals and the pattern of freezing in peach flower buds. In dormant buds, ice crystals formed at localized sites within the bud axis and scales. Ice crystal formation disrupted tissues and mechanical injury from repetitive freezethaw cycles was apparent. There was evidence of ice formation in the floral organs of dormant buds exposed to ?25°C but none observed in buds exposed to either ?5 or ?10°C. The distribution of ice crystals was different in deacclimated buds. In addition to large ice crystals within the subtending bud axis and scales, evidence of large crystals within the developing floral organs was noted. These crystals were most prominent in the lower portions of the developing flower and peduncle, and caused a separation of the epidermal layer from adjacent cells. The distribution of ice crystals within both dormant and deacclimated peach flower buds corroborated the results of previous thermal analysis experiments.     

Frost Injury as a Possible Inciting Factor in Bud and Shoot Necroses of Fraxinus excelsior L.

Large numbers of European ash have died in Poland in all age classes during the last ten years. The characteristic symptom occurring on shoots of planted and self‐sown seedlings was bark necroses starting from the shoot apex, necrotic buds, or leaf and twig scars. The results showed that in the bud tissue of cold acclimated European ash extracellular and intracellular ice formation occurred at approximately ?9 and ?32°C, respectively. In deacclimated plants in spring water supercooling is limited by the heterogenous ice nucleation temperature and consequently the cold tolerance is ?9 to ?4°C for bud tissues and ?13 to ?9°C for shoots. Isolations of fungi were performed from dead buds and from necroses occurring on the main stem. Alternaria alternata, Fusarium lateritium and Phomopsis scobina were among the fungi occurring in both these organs at frequencies of more than 7%. Cylindrocarpon heteronemum, Diplodia mutila and Tubercularia vulgaris from necroses were only isolated in frequencies; 3.3, 1.2 and 5.4%, respectively. It seems likely that freezing injury is the inciting factor, which combined with fungal colonization manifests itself as fatal damage to European ash buds and shoots.     

Effects of Temperature on Cold Acclimation and Deacclimation in Flower Buds of Evergreen Azaleas

The effects of various storage temperature/duration combinations(5, 10 and 17°/4, 8, 12 and 16 weeks) on cold acclimationand deacclimation of flower buds were studied in four speciesof evergreen azaleas having different natural distribution andcold hardiness. The freezing process and the exotherm temperaturedistribution of florets in excised whole buds determined bydifferential thermal analysis were used as the diagnostics todetermine the degree of bud acclimation and deacclimation. Theacclimation in buds lasted for as long as 12 to 16 weeks at5°C storage, and from 8 to 12 weeks at 10°C, and itappeared to be maintained after the chilling requirement forbreaking bud dormancy had been satisfied. Therefore, bud acclimationseems to be maintained independently from bud dormancy. Thedehardening effect on acclimated buds occurred as a result ofshort exposures to higher temperatures or long exposures tolower temperatures, and there was no relation between the rateof deacclimation and the degree of hardiness in each species.Among three storage temperatures examined, 5°C was the mosteffective for the maintenance of cold acclimation in flowerbuds and the small difference of floret water contents at 5and 10°C storage is not significant. (Received August 28, 1982; Accepted February 4, 1983)     

Effects of prescribed burning on leaves and flowering of Quercus garryana

Many woodland understories are managed with prescribed fire. While prescribed burns intended to manipulate understory vegetation and fuels usually do not cause excessive tree mortality, sublethal canopy damage may occur and can affect tree vigor and reproductive output. We monitored Quercus garryana trees in western Washington, USA with multiple canopy thermocouples during three prescribed burns. Peak temperatures recorded in tree canopies ranged from 36 to 649°C. We assessed leaf damage immediately after burning, and flower, leaf and acorn production in the following year in the vicinity of each thermocouple. Leaf scorch first occurred with peak thermocouple temperatures around 45°C, was variable up to 75°C, but above 75°C all leaves were killed. Buds, including their reproductive and leaf organs were more resistant to heat damage than leaves, but leaf scorch had predictive value in forecasting bud organ damage. Staminate and pistillate inflorescences and acorn production per bud decreased and bud mortality increased with maximum thermocouple temperature. In two burns where the highest peak temperatures reached 137°C, there was no difference in leaf production between burned and control plots in the spring following burning. However, no staminate or pistillate inflorescences were produced when thermocouple peak temperatures went above 55 or 68°C, respectively. While heat damage to bud organs was detected, production of reproductive organs was also curtailed at temperatures lower than could reasonably be attributed to heat damage. Thus, it is probable that some other fire-related factor, possibly smoke, was also involved.     

Freezing behaviours in wintering Cornus florida flower bud tissues revisited using MRI

How plant tissues control their water behaviours (phase and movement) under subfreezing temperatures through adaptative strategies (freezing behaviours) is important for their survival. However, the fine details of freezing behaviours in complex organs and their regulation mechanisms are poorly understood, and non‐invasive visualization/analysis is required. The localization/density of unfrozen water in wintering Cornus florida flower buds at subfreezing temperatures was visualized with high‐resolution magnetic resonance imaging (MRI). This allowed tissue‐specific freezing behaviours to be determined. MRI images revealed that individual anthers and ovules remained stably supercooled to ?14 to ?21 °C or lower. The signal from other floral tissues decreased during cooling to ?7 °C, which likely indicates their extracellular freezing. Microscopic observation and differential thermal analyses revealed that the abrupt breakdown of supercooled individual ovules and anthers resulted in their all‐or‐nothing type of injuries. The distribution of ice nucleation activity in flower buds determined using a test tube‐based assay corroborated which tissues primarily froze. MRI is a powerful tool for non‐invasively visualizing unfrozen tissues. Freezing events and/or dehydration events can be located by digital comparison of MRI images acquired at different temperatures. Only anthers and ovules preferentially remaining unfrozen are a novel freezing behaviour in flower buds. Physicochemical and biological mechanisms/implications are discussed.     

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.     

Winter bud content according to position in 3-year-old branching systems of 'Granny Smith' apple

An investigation was made of the number of preformed organs in winter buds of 3-year-old reiterated complexes of the 'Granny Smith' cultivar. Winter bud content was studied with respect to bud position: terminal buds were compared on both long shoots and spurs according to branching order and shoot age, while axillary buds were compared between three zones (distal, median and proximal) along 1-year-old annual shoots in order 1. The percentage of winter buds that differentiated into inflorescences was determined and the flowers in each bud were counted for each bud category. The other organ categories considered were scales and leaf primordia. The results confirmed that a certain number of organs must be initiated before floral differentiation occurred. The minimum limit was estimated at about 15 organs on average, including scales. Total number of lateral organs formed was shown to vary with both bud position and meristem age, increasing from newly formed meristems to 1- and 2-year-old meristems on different shoot types. These differences in bud organogenesis depending on bud position, were consistent with the morphogenetic gradients observed in apple tree architecture. Axillary buds did not contain more than 15 organs on average and this low organogenetic activity of the meristems was related to a low number of flowers per bud. In contrast, the other bud categories contained more than 15 differentiated organs on average and a trade-off was observed between leaf and flower primordia. The ratio between the number of leaf and flower primordia per bud varied with shoot type. When the terminal buds on long shoots and spurs were compared, those on long shoots showed more flowers and a higher ratio of leaf to flower primordia.     

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.     

Similarity in the effects of different seed cryoconservation conditions on plant growth and development

Morphological traits of 35 wild plant species were studied after freezing the seeds using quick or programmed two-stage freezing to ?196°C and nondeep freezing to ?10°C. The seeds were stored frozen for 1 month. The seeds of all studied species had low moisture content. Field and laboratory germination rates, leaf growth, the number and length of shoots, the number of generative organs, and variation of all these traits were studied. It was demonstrated that the traits changed in the same directions independently of the freezing conditions, except for laboratory germination rate for some species. The changes were determined by the species-specific plant characteristics rather than by the freezing mode.     

Bud damage from controlled heat treatments in Quercus garryana

Quercus garryana habitats are increasingly being managed with prescribed fire, but acorn dependent wildlife might be adversely affected if fires damage acorn crops. We examined one way that fire might affect subsequent acorn crops: through direct heating and damage of buds containing the following year’s floral organs. We measured internal bud temperatures during controlled time and temperature treatments, described damage to heated buds at the tissue and cellular levels and quantified spring flowering to assess the consequences of the treatments. We found that internal bud temperature was logarithmically related to exposure time and linearly related to treatment temperature. Tissue damage was more common in bud scales, staminate and bud scale scar primordia than in leaf, pistillate, leaf axillary primordia and apical meristems. Damaged tissues were sequestered by cells with thickened cell walls. A 133°C treatment applied for 60 s produced minimal damage or mortality, but damage increased rapidly in hotter or longer treatments, culminating in 100% mortality at 273°C for 60 s. Our experiments account only for radiative, not convective heating, but suggest that fires might produce sublethal effects that affect flowering and acorn crops. Q. garryana’s large buds possess an internal organ arrangement well suited to minimizing heat damage. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The Effect of Temperature on the Production and Abscission of Flowers and Pods in Snap Bean (Phaseolus vulgaris L.)

The effect of temperature on production and abscission of flowerbuds, flowers and pods was studied in a determinate snap-beancultivar (cv. Tenderette). Under moderate temperature (e.g.27/17 °C) the onset of pod development was associated withcessation of flower bud production and with enhanced abscissionof flower buds. Raising night temperature from 17 °C to27 °C strongly reduced pod production, mature pod size andseeds per pod, while an increase in day temperature from 22°C to 32 °C had smaller and less consistent effects.Pod production under high night temperature was not constrainedby flower production since 27 °C at night promoted branchingand flower bud appearance. Under 32/27 °C day/night temperaturethe large reduction in pod set was due to enhanced abscissionof flower buds, flowers and young pods (< 3 cm). Flowershad the highest relative abscission followed by young pods andflower buds. Therefore, the onset of anthesis and of pod developmentwere the plant stages most sensitive to night temperature. Podslarger than 3 cm did not abscise but usually aborted and shrivelledunder high night temperature. The effects of 32/27 °C werenot due to transient water stresses and were observed even undercontinuous irrigation and mist-spraying. High temperature, flower production, pod set, seed set, abscission, snap bean, Phaseolus vulgaris L., cv. Tenderette     

Latitudinal variation in suppression of flower bud formation at high temperature in four native dandelions (Taraxacum spp.) with different distributions and genetic structures in Japan

The control of the response of flowering to temperature plays a key role in successful range‐expansion of plants. A previous study showed that the suppression of flower‐bud formation at high temperature in Taraxacum officinale decreases genetically with latitude from north to south in Japan. The present study investigated whether similar trait variation occurs among populations of native Taraxacum species in Japan. Seedlings of T. albidum (a low‐ and mid‐latitude allopolyploid), T. japonicum (a mid‐latitude diploid) and T. venustum (a high‐latitude autopolyploid) were grown at three temperatures. Time to flower‐bud appearance increased with temperature in T. japonicum and T. venustum, but did not increase in T. albidum. Time to flower‐bud appearance did not differ significantly among the three species at 14°C, but it was shorter in T. albidum than in the other two species at 19°C and 24°C. The early appearance of buds of T. albidum was confirmed by another experiment in which plants of 18 populations from the three species and T. platycarpum (a mid‐latitude diploid) grown at 19°C were used. The results clearly indicate that high‐temperature suppression of flower‐bud formation was lower in low‐latitude species than in high‐latitude species. This interspecific variation is analogous to the intraspecific variation in T. officinale. Time to bud appearance of five populations in T. albidum was homogeneous within and between the populations. The results suggest that the five populations are monoclonal and lack the sensitivity of suppression of flower‐bud formation to high temperature.     

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.     

越冬代梨木虱、梨二叉蚜产卵选择对子代种间竞争的影响

梨树萌芽期,越冬代梨木虱Cacopsylla chinensis和梨二叉蚜Schizaphis piricola子代共用同一食物资源,为了解其母代如何避免子代发生种间竞争,通过研究其产卵特点表明,梨二叉蚜和梨木虱在梨树枝条上的产卵位点不同。梨木虱主要在花芽枝上进行产卵,偏好于花芽芽鳞和枝条基部刻痕处产卵,以枝条基部产卵量最多,其次为顶花芽,产卵量随芽位次序依次下降,梨木虱还可在叶芽部位产卵,但数量较少;梨二叉蚜主要于叶芽枝上进行产卵,偏好在叶芽枝的第2、3芽位产卵,顶芽和枝条基部无产卵,未见梨二叉蚜在花芽部位产卵;在花芽枝上,梨木虱卵的空间生态位大于梨二叉蚜、叶芽枝上却相反,两者在梨树枝条上的产卵空间生态位呈分离状态;梨木虱和梨二叉蚜孵化的若虫数量及空间分布特点与卵一致;对梨木虱和梨二叉蚜产卵位点分析表明,其产卵位点的物理性状明显不同。本研究明确了梨木虱和梨二叉蚜母代通过不同产卵位点的选择,降低了子代种间的同位竞争。     

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.     

Supercooling Ability, Water Content and Hardiness of Rhododendron Flower Buds during Cold Acclimation and Deacclimation

The relationship between supercooling ability and water contentand killing temperature of flower buds during cold acclimationand deacclimation were studied using R. kiusianum and R. x akebono.The occurrence of multiple floret exotherms and their shiftto a narrow range at lower subzero temperatures, as well asthe marked decrease of florets water content, were observedas the symptoms of cold acclimation occuring in flower budsfrom fall to winter, and vice versa in spring buds during deacclimation.In R. kiusianum, the fully acclimated period was from Novemberto March and two months longer than that of R. x akebono. Thesupercooling ability of the former was about –25°Cand about –20°C in the latter. Although the watermigration within bud tissues during the freezing process wasdetermined in the acclimated and deacclimated buds for R. xakebono, no significant water changes could be observed, evenin the acclimated buds. Thus, it is conceivable that deep supercoolingin florets may result not necessarily from water migration fromflorets and bud axes to scales in response to freezing, butfrom low water content in situ of cold-acclimated or artificiallydehydrated flower buds. (Received July 29, 1981; Accepted October 12, 1981)     

Partitioning of [14C]sucrose and Acid Invertase Activity in Reproductive Organs of Pepper Plants in Relation to Their Abscission Under Heat Stress

The effect of heat stress on processes related to carbohydratepartitioning was investigated in young bell pepper (Capsicumannum L. cv. Maor) plants in relation to abscission of theirreproductive organs at different stages of development. None of the reproductive organs abscised after 5 d in a normalday/night temperature regime (25/18 °C). With a temperatureregime of 35 °C day, 25 °C night, abscission occurredin only a small portion of the flower buds and none of the flowersand fruitlets. However, when temperatures in the day and nightwere reversed (25/35 °C, day/night) all the buds and someof the flowers abscised during that time period. The young fruitat the first node did not abscise under any temperature regime.The abscission rate of the flower buds was reduced under heatstress if the developing fruit at the first node had been removed. High temperature during either the light or dark periods reducedthe export of [¹⁴C]sucrose from the source leaf (fed for 48h with [¹⁴C]sucrose). Both heat stress and fruit presence reduced the relative amountof [¹⁴C]sucrose which was exported to the flower buds, flowersand roots. Likewise, these treatments reduced the concentrationof reducing sugars in the reproductive organs. Concomitantly,the heat stress and fruit presence on the first node reducedthe activity of soluble acid invertase in the flower buds andthe roots, but not in young leaves. Overall, the results show that heat stress causes alternationin sucrose distribution in the plant, but may also have specificeffects on metabolic activities related to sucrose import andutilization in flower buds and flowers which in turn may enhancetheir abscission. Bell pepper, (Capsicum annuum L. cv. Maor), abscission, acid invertase, heat stress, reproductive organs, sink leaves     

Freezing Tolerance of Shoot and Flower Primordia of Coniferous Buds by Extraorgan Freezing

Shoot and flower primordia of vegetative and flower buds ofextremely or very hardy conifers belonging to the subfamilyAbietoideae of the Pinaceae, survived between –40 and–70?C by extraorgan freezing, which differed greatly dependingupon species. The water in these organs gradually froze outwith decreasing temperatures when cooled very slowly, whichenabled these organs to survive %40?C or below. The same icesegregation in shoot and flower primordia by extraorgan freezingwas observed in most of the temperate conifers belonging toTaxaceae, Cephalotaxaceae, Taxodiaceae and Cuppressaceae, makingthem resistant to temperatures between –15 and –25?C.In these conifers, scales acted as an ice sink, unlike the conifersof Abietoideae. The rates of cooling and exosmosis of waterin the shoot or flower primordia, their size, and their abilityto tolerate freeze-dehydration or its related stress play animportant role in determining whether death is caused by freeze-dehydrationor intraorgan freezing. Even in very hardy conifers, low temperature exotherms fromfreezing within the shoot primordia appeared between –30and –35?C on the DTA profiles when cooled continuouslyunder laboratory conditions from 5?C to –50?C at 2 to5?C/h. Appearance of low temperature exotherms always resultedin death. However, in the coldest area of Hokkaido, where theair temperature cools down to –40?C or below nearly everyyear, such an intraorgan freezing seems seldom to occur, especiallyin natural stands. On the other hand, low temperatures below–25?C seldom occur in warm-temperate climates. Thus, itmay be considered that in both boreal and temperate coniferstheir shoot and flower primordia seem to tolerate freeze dehydrationby extraorgan freezing under natural conditions. ¹ Contribution No. 2431 from the rnstitute of Low TemperatureScience. (Received March 27, 1982; Accepted August 12, 1982)     

Winter climate change and the poleward range expansion of a tropical invasive tree (Brazilian pepper—Schinus terebinthifolius)

Winter climate change is expected to lead to the tropicalization of temperate ecosystems, where tropical species expand poleward in response to a decrease in the intensity and duration of winter temperature extremes (i.e., freeze events). In the southeastern United States, freezing temperatures control the northern range limits of many invasive nonnative species. Here, we examine the influence of freezing temperatures and winter climate change on the northern range limits of an invasive nonnative tree—Schinus terebinthifolius (Brazilian pepper). Since introduction in the 1800s, Brazilian pepper has invaded ecosystems throughout south and central Florida to become the state's most widespread nonnative plant species. Although Brazilian pepper is sensitive to freezing temperatures, temperature controls on its northern distribution have not been adequately quantified. We used temperature and plant occurrence data to quantify the sensitivity of Brazilian pepper to freezing temperatures. Then, we examined the potential for range expansion under three alternative future climate scenarios (+2°C, +4°C, and +6°C). Our analyses identify a strong nonlinear sigmoidal relationship between minimum temperature and Brazilian pepper presence, with a discrete threshold temperature occurring near ?11°C. Our future scenario analyses indicate that, in response to warming winter temperatures, Brazilian pepper is expected to expand northward and transform ecosystems in north Florida and across much of the Gulf of Mexico and south Atlantic coasts of the United States. These results underscore the importance of early detection and rapid response efforts to identify and manage the northward invasion of Brazilian pepper in response to climate change. Looking more broadly, our work highlights the need to anticipate and prepare for the tropicalization of temperate ecosystems by tropical invasive species.     

The Formation and Distribution of Ice within Forsythia Flower Buds

Differential thermal analysis detected two freezing events when dormant forsythia (Forsythia viridissima Lindl.) flower buds were cooled. The first occurred just below 0°C, and was coincident with the freezing of adjacent woody tissues. The second exotherm appeared as a spike between −10 and −25°C and was correlated with the lethal low temperature. Although this pattern of freezing was similar to that observed in other woody species, differences were noted. Both direct observations of frozen buds and examination of buds freeze-fixed at −5°C demonstrated that ice formed within the developing flowers at temperatures above the second exotherm and lethal temperature. Ice crystals had formed within the peduncle and in the lower portions of the developing flower. Ice also formed within the scales. In forsythia buds, the developing floral organ did not freeze as a unit as noted in other species. Instead the low temperature exotherm appeared to correspond to the lethal freezing of supercooled water within the anthers and portions of the pistil.