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Cindy L. Flinn and Edward N. Ashworth

Examination of both frozen specimens and -5C freeze-fixed buds showed that ice crystals were not uniformly distributed in blueberry flower buds. Localized freezing was also evidenced by detection of multiple freezing events using differential thermal analysis (DTA). Upon cooling, an initial exotherm occurred just below 0C and coincided with ice formation in adjacent woody tissue. Multiple low temperature exotherms (LTE), which have been reported to correspond with the freezing of individual blueberry florets (Bierman, et al. 1979. ASHS, 104(4):444-449), occurred between -7C and -28C. The presence and temperature of LTEs was influenced by cooling rates and whether buds were excised. LTE temperatures did not correlate with hardiness of buds frozen under field-like conditions. Results suggested that DTA of excised buds was not an appropriate method for determining hardiness.

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John Carter, Rex Brennan, and Michael Wisniewski

Ice formation and movement in stems, leaves, and flowers of blackcurrant were observed by infrared video thermography. Stem sections bearing leaves and racemes were cooled slowly to as low as -6.4 °C and allowed to freeze without artificial nucleation. Ice formed in stems first, then moved from stems into leaves and racemes. Patterns of ice movement were complex and depended upon the temperature of the initial nucleation event. Individual flowers froze between -1.6 and -5.5 °C. Survival of flowers after a cooling treatment depended upon whether they froze and the amount of freezing that occurred in the peduncles to which they were attached. Some flowers survived the initial freezing treatments but later died because of peduncle damage. Movement of ice from stems into peduncles sometimes was observed to occur in discrete steps, separated by time and temperature. Several independent freezing events were often observed in a peduncle, rather than one continuous event. Pedicels attached to frozen peduncles often remained supercooled for several minutes to over an hour before freezing. No consistent pattern was evident during freezing of individual flowers in an inflorescence. The range of temperature over which flowers in a single inflorescence froze was in some instances over 4 °C. Both mature and immature flowers supercooled. Barriers to movement of ice appeared to exist at certain anatomical junctions within the plant, notably where the peduncle of an inflorescence attaches to a stem and where a flower pedicel joins a peduncle. The time required for ice to pass through these barriers was inversely related to the degree of supercooling that had occurred prior to freezing.

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Michele R. Warmund, Fumiomi Takeda, and Glen A. Davis

`Hull Thornless' and `Black Satin' blackberry (Rubus spp.) canes were collected from Sept. 1989 through Mar. 1990 to determine the hardiness and supercooling characteristics of buds at various stages of development. Anatomical studies were also conducted to examine the location of ice voids in buds frozen to -5 or -30C. Differentiation of the terminal flower occurred in `Black Satin' buds by 6 Nov., whereas `Hull Thornless' buds remained vegetative until early spring. As many as nine floral primordia were observed in both cultivars by 12 Mar. The hardiness of the two cultivars was similar until February. Thereafter, `Black Satin' buds were more susceptible to cold injury than those of `Hull Thornless'. Flora1 and undifferentiated buds of both cultivars exhibited one to four low temperature exotherms (LTEs) from 9 Oct. to 12 Mar. in differential thermal analysis (DTA) experiments. The stage of flora1 development did not influence the bud's capacity to supercool. The number of LTEs was not related to the stage of floral development or to the number of floral primordia. Extracellular voids resulting from ice formation in the bud axis and scales were observed in samples subjected to -5 or -30C.

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Michele R. Warmund, Rusty Fuller, and John H. Dunn

Rhizomes of `Meyer' zoysiagrass (Zoysia japonica Steud.) were subjected to temperatures below 0 °C and were subsequently placed in a growth chamber with air at 34 °C day/28 °C night to determine the rate of shoot growth from nodes. Rhizomes exposed to subzero temperatures produced shoots steadily up to 16 days after freezing (DAF), but subsequent shoot growth from rhizomes was minimal. At 32 DAF, shoots were present on 68% and 44% of the nodes of unfrozen control (2 °C) rhizomes and those frozen to -7 °C, respectively. In another study, samples were frozen to a sublethal temperature (-7 °C) to examine the distribution of extracellular ice voids near the apical meristems of rhizomes and to characterize tissue recovery. Extracellular voids were present within the leaf tissue and between the leaves in samples prepared for scanning electron microscopy (SEM) immediately after freezing to -7 °C. By 12 DAF, most of the remaining voids were observed in older leaves. Nearly all extracellular voids in the leaves were absent by 20 DAF. However, by 28 DAF, some rhizomes still had small voids between leaves. Although the structure of zoysiagrass rhizomes subjected to -7 °C was temporarily disrupted, tissues recovered from extracellular freezing and new shoot growth was produced following exposure to warm temperatures.

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Beth Ann A. Workmaster, Jiwan P. Palta, and Michael Wisniewski

Infrared video thermography was used to study formation of ice in leaves, stems, and fruit of cranberry (Vaccinium macrocarpon Ait. `Stevens'). Ice formed on the plant surface at -1 or -2 °C by freezing of a droplet of water containing ice nucleation-active bacteria (Pseudomonas syringae van Hall). Samples were then cooled to a minimum of -8 °C. Observations on the initiation and propagation of ice were recorded. Leaves froze only when ice was present on the abaxial surface. Once initiated, ice propagated to the stem and then readily to other leaves. In both unripe and ripe fruit, ice propagation from the stem to the fruit via the pedicel was not observed. Fruit remained supercooled for up to 1 hour after ice was present in the stem. Fruit could only be nucleated when ice was present at the calyx (distal) end. Red (ripe) berries supercooled to colder temperatures and for longer durations than blush (unripe) berries before an apparent intrinsic nucleation event occurred. These observations provide evidence that leaves are nucleated by ice penetration via stomata. The ability of fruit to supercool appears to be related to the presence of barriers to extrinsic ice propagation at both the pedicel and fruit surface. Stomata at the calyx end of the fruit in the remnant nectary area may provide avenues for extrinsic ice nucleation.

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Michael Wisniewski

Frost-sensitive plant species have a limited ability to tolerate ice formation in their tissues. Most plants can supercool below 0°C and avoid ice formation. Discrepancies exist about the role of intrinsic and extrinsic ice-nucleating agents in initiating ice formation in plants. Previous research has demonstrated the ability of infrared video thermography to directly observe and record the freezing process in plants (Wisniewski et al., 1997. Plant Physiol. 113:4378–4397). In the present study, the ability of droplets of a suspension of the ice-nucleating-active (Ice+) bacterium, Pseudomonas syringae, and droplets of deionized water, to induce ice formation in bean plants was compared. The activity of these agents were also compared to intrinsic ice formation in dry plants. Results indicated that the presence of the Ice+ bacteria in droplets ranging from 0.5–4.0 μL always induced freezing at a warmer temperature than droplets of deionized water alone (no bacteria) or intrinsic nucleators in dry plants. When droplets of Ice+ bacteria were allowed to dry, they were no longer effective but were active again upon rewetting. Droplets of water would often supercool below temperatures at which ice formation was initiated by intrinsic agents. When a silicon grease barrier was placed between the droplets of Ice+ bacteria and the leaf surface, the bacteria were no longer capable of inducing ice formation in the plant, despite the droplets being frozen on the plant surface. This indicates that ice crystals must penetrate the cuticle in order to induce freezing of the plant.

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Steven E. Lindow

Genes determining the ability of the bacterium Pseudomonas syringae to catalyze ice formation have been cloned and characterized. Ice nucleation active (Ice+) strains of this species are common on plants and the supercooling ability of frost sensitive plants is inversely proportional to the logarithm of the population size of Ice+ bacteria at temperatures above -5C. Recombinant Ice- derivatives off. syringae were produced by site-directed mutagenesis using deletion containing ice genes cloned form this species. The Ice- strains colonized potatoes well in field studies, reduced the population size of Ice+ bacterial strains by about 50-fold, and reduced the incidence of frost injury an average of 82% in several radiative frosts of temperatures in the range of -3 to -5 C. The ice gene has also been introduced into Solanum commersonii to determine its effect on increasing the tolerance of ice formation in this frost tolerant species. Transgenic plants exhibit a much higher threshold ice nucleation temperature than the parental plants.

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R.C. Ebel, P.A. Carter, W.A. Dozier, D.A. Findley, M.L. Nesbitt, B.R. Hockema, and J.L. Sibley

The current study was conducted to relate ice formation to the pattern and rate of leaf and stem injury of Satsuma mandarins on trifoliate orange rootstock. Potted trees were unacclimated, moderately acclimated or fully acclimated by exposing trees to 32/21 °C, 15/7 °C or 10/4 °C, respectively. Freezing treatments consisted of decreasing air temperature at 2 °C·h-1 until ice formed as evidenced by exotherms determined using differential thermal analysis of stems. Air temperature was then decreased, held constant, or increased and held constant to simulate severe, moderate and mild freeze conditions, respectively. All treatment exhibited exotherms at -2 to -4 °C, which were smaller with milder freezing treatments. Only the fully acclimated trees exhibited multiple exotherms. Leaf watersoaking, an indication of ice formation, occurred concurrently with stem exotherms except for fully acclimated trees where there was up to a 30-min delay and which corresponded with the second exotherm. Electrolyte leakage of leaves began to increase near the peak of the stem exotherm, but increased more slowly with milder freezing temperature treatments. In some treatments, electrolyte leakage reached a plateau near 50% but leaves survived. Leaves died when whole-leaf electrolyte leakage exceeded 50%. These data are discussed within the framework of a proposed mechanism of injury of Satsuma mandarin leaves by subfreezing temperatures, especially multiple exotherms of fully acclimated trees, and the plateau of electrolyte leakage of leaves at the critical level for survival.

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Cindy L. Flinn and Edward N. Ashworth

The location of ice crystals and their relationship to xylem vessels was studied in nonacclimated and acclimated `Berkeley' blueberry (Vaccinium corymbosum L.) flower buds. Light microscopy and low-temperature scanning electron microscopy (SEM) were used to detect ice crystals in the bud scales, floret scales, and bracts of dormant flower buds that had been frozen to -15C. No evidence of ice formation was observed in rachises, pedicels, and organs in florets when buds that had been fixed while frozen at -5C were examined with conventional SEM. This indicated that dormant buds underwent extraorgan freezing as a survival mechanism. Ice formation was not uniform in nonacclimated or deacclimated buds, although it was more prevalent in both than in acclimated buds. Large ice crystals were found in the ovaries of freeze-stressed nonacclimated buds. In deacclimated freeze-stressed buds, ice was found in the petals, rachises, pedicels, and ovaries. To determine whether this ice distribution pattern was correlated with the presence of mature xylem vessels, cleared flower buds were stained with basic fuchsin, which revealed the intact network of lignified elements. In nonacclimated buds (20 Sept.), mature xylem vessels extended through the rachises, connecting the bud scales with the floret scales and through the pedicels into the corollas of the florets. Although vascular development occurred in dormant buds, the greatest proliferation of vessels in the ovaries, petals, and sepals occurred coincident to the appearance of ice in these organs and the loss of hardiness.

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M.A. McKellar, D.W. Buchanan, Dewayne L. Ingram, and C.W. Campbell

Freezing tolerance and the lethal freezing temperature were determined for detached leaves of avocado (Persea americana Mill.) by either electrolyte leakage or visual appearance of browning. Leaves from field-grown trees of `Gainesville', `Booth8', and `Winter Mexican' in both Gainesville and Homestead, Fla., were evaluated. All cultivars in both locations survived ice formation in their tissue. Leaf tissue had a temperature limit (lethal freeze temperature) at and below which the tissue died. The lethal freezing temperature varied from -5.1 to -9.3C, depending on time of year and location. The lethal freeze temperature for a cultivar decreased over the fall and winter as temperatures decreased. Leaves of `Booth-8' and Winter Mexican' decreased 2.5 and 1.5C, respectively, in Homestead from 13 Nov. 1982 to 5 Feb. 1983. The plants growing at the lower temperature location (Gainesville) had lower lethal freeze temperatures. Leaves of `Gainesville' had a lethal freeze temperature of - 9.3C from trees at Gainesville and - 7.8C from trees at Homestead.