细菌冰核提高印度谷螟过冷却点的研究

印度谷螟(Plodia interpunctella)是一种不耐结冰的昆虫,在冬季它通过降低过冷却 点以避免结冰。现已查明,冰核活性细菌能显著提高植物的过冷却点,导致许多作物在较高 的温度下发生霜冻害。本文也证明细菌冰核能显著提高印度谷螟虫的过冷却点。对照的平均过冷却点是-17.6℃;分别用0.1g和1g细菌冰核与1kg面粉混合后进行处理,平均过冷却点分别比对照提高了12.8℃和13.6℃。研究结果支持这样的观点：细菌冰核有可能成为一种在冬季使用的、杀灭不耐结冰害虫的生物制剂。     

Change of supercooling capability in solutions containing different kinds of ice nucleators by flavonol glycosides from deep supercooling xylem parenchyma cells in trees

Deep supercooling xylem parenchyma cells (XPCs) in Katsura tree contain flavonol glycosides with high supercooling-facilitating capability in solutions containing the ice nucleation bacterium (INB) Erwinia ananas, which is thought to have an important role in deep supercooling of XPCs. The present study, in order to further clarify the roles of these flavonol glycosides in deep supercooling of XPCs, the effects of these supercooling-facilitating (anti-ice nucleating) flavonol glycosides, kaempferol 3-O-β-d-glucopyranoside (K3Glc), kaempferol 7-O-β-d-glucopyranoside (K7Glc) and quercetin 3-O-β-d-glucopyranoside (Q3Glc), in buffered Milli-Q water (BMQW) containing different kinds of ice nucleators, including INB Xanthomonas campestris, silver iodide and phloroglucinol, were examined by a droplet freezing assay. The results showed that all of the flavonol glycosides promoted supercooling in all solutions containing different kinds of ice nucleators, although the magnitudes of supercooling capability of each flavonol glycoside changed in solutions containing different kinds of ice nucleators. On the other hand, these flavonol glycosides exhibited complicated nucleating reactions in BMQW, which did not contain identified ice nucleators but contained only unidentified airborne impurities. Q3Glc exhibited both supercooling-facilitating and ice nucleating capabilities depending on the concentrations in such water. Both K3Glc and K7Glc exhibited only ice nucleation capability in such water. It was also shown by an emulsion freezing assay in BMQW that K3Glc and Q3Glc had no effect on homogeneous ice nucleation temperature, whereas K7Glc increased ice nucleation temperature. The results indicated that each flavonol glycoside affected ice nucleation by very complicated and varied reactions. More studies are necessary to determine the exact roles of these flavonol glycosides in deep supercooling of XPCs in which unidentified heterogeneous ice nucleators may exist.     

High ice nucleating ability in plant leaves

A high ice nucleating ability in leaves of Veronica persicaand Buxus microphylla was observed on analyzing the frequencydistribution of their freezing temperatures. The most efficientice nucleators in Veronica leaves exist in structures withinthe leaf blades (mesophyll), and those of Buxus in structureswithin the midrib (vascular tissue). Dissolved or suspendedleaf structures were much less effective. (Received March 16, 1973; )     

The inhibition of ice nucleators by insect antifreeze proteins is enhanced by glycerol and citrate

Antifreeze proteins depress the freezing point of water while not affecting the melting point, producing a characteristic difference in freezing and melting points termed thermal hysteresis. Larvae of the beetle Dendroides canadensis accumulate potent antifreeze proteins (DAFPs) in their hemolymph and gut, but to achieve high levels of thermal hysteresis requires enhancers, such as glycerol. DAFPs have previously been shown to inhibit the activity of bacterial and hemolymph protein ice nucleators, however, the effect was not large and therefore the effectiveness of the DAFPs in promoting supercooling of the larvae in winter was doubtful. However, this study demonstrates that DAFPs, in combination with the thermal hysteresis enhancers glycerol (1 M) or citrate (0.5 M), eliminated the activity of hemolymph protein ice nucleators and Pseudomonas syringae ice-nucleating active bacteria, and lowered the supercooling points (nucleation temperatures) of aqueous solutions containing these ice nucleators to those of water or buffer alone. This shows that the DAFPs, along with glycerol, play a critical role in promoting hemolymph supercooling in overwintering D. canadensis. Also, DAFPs in combination with enhancers may be useful in applications which require inhibition of ice nucleators.     

The Freezing Response of an Arctic Cushion Plant, Saxifraga caespitosa L.: Acclimation, Freezing Tolerance and Ice Nucleation

Cold hardiness in actively growing plants of Saxifraga caespitosaL., an arctic and subarctic cushion plant, was examined. Plantscollected from subarctic and arctic sites were cultivated ina phytotron at temperatures of 3, 9, 12 and 21 °C undera 24-h photoperiod, and examined for freezing tolerance usingcontrolled freezing at a cooling rate of 3–4 °C eitherin air or in moist sand. Post-freezing injury was assessed byvisual inspection and with chlorophyll fluorescence, which appearedto be well suited for the evaluation of injury in Saxifragaleaves. Freezing of excised leaves in moist sand distinguishedwell among the various treatments, but the differences werepartly masked by significant supercooling when the tissue wasfrozen in air. Excised leaves, meristems, stem tissue and flowerssupercooled to –9 to –15 °C, but in rosettesand in intact plants ice nucleation was initiated at –4to –7 °C. The arctic plants tended to be more coldhardy than the subarctic plants, but in plants from both locationscold hardiness increased significantly with decreasing growthtemperature. Plants grown at 12 °C or less developed resistanceto freezing, and excised leaves of arctic Saxifraga grown at3 °C survived temperatures down to about –20 °C.Exposure to –3 °C temperature for up to 5 d did notsignificantly enhance the hardiness obtained at 3 °C. Whenwhole plants of arctic Saxifraga were frozen, with roots protectedfrom freezing, they survived –15 °C and –25°C when cultivated at 12 and 3 °C, respectively, althougha high percentage of the leaves were killed. The basal levelof freezing tolerance maintained in these plants throughoutperiods of active growth may have adaptive significance in subarcticand arctic environments. Saxifraga caespitosa L., arctic, chlorophyll fluorescence, cold acclimation, cushion plant, freezing stress, freezing tolerance, ice nucleation, supercooling     

Freezing avoidance by supercooling in Olea europaea cultivars: the role of apoplastic water,solute content and cell wall rigidity

Plants can avoid freezing damage by preventing extracellular ice formation below the equilibrium freezing temperature (supercooling). We used Olea europaea cultivars to assess which traits contribute to avoid ice nucleation at sub‐zero temperatures. Seasonal leaf water relations, non‐structural carbohydrates, nitrogen and tissue damage and ice nucleation temperatures in different plant parts were determined in five cultivars growing in the Patagonian cold desert. Ice seeding in roots occurred at higher temperatures than in stems and leaves. Leaves of cold acclimated cultivars supercooled down to ?13 °C, substantially lower than the minimum air temperatures observed in the study site. During winter, leaf ice nucleation and leaf freezing damage (LT₅₀) occurred at similar temperatures, typical of plant tissues that supercool. Higher leaf density and cell wall rigidity were observed during winter, consistent with a substantial acclimation to sub‐zero temperatures. Larger supercooling capacity and lower LT₅₀ were observed in cold‐acclimated cultivars with higher osmotically active solute content, higher tissue elastic adjustments and lower apoplastic water. Irreversible leaf damage was only observed in laboratory experiments at very low temperatures, but not in the field. A comparative analysis of closely related plants avoids phylogenetic independence bias in a comparative study of adaptations to survive low temperatures.     

Supercooling along an Altitudinal Gradient in Espeletia schultzii, a Caulescent Giant Rosette Species

Rada, F., Goldstein, G., Azocar, A. and Torres, F. 1987. Supercoolingalong an altitudinal gradient in Espeletia schultzii, a caulescentgiant rosette species.—J. exp. Bot. 38: 491–497. Tropical high Andes plants may be exposed to sub-zero temperaturesany night of the year. These plants have to rely on mechanismswhich protect them from these environmental conditions but atthe same time allow their growth and development. Supercoolinghas been found to be the principal avoidance mechanism in leavesof the caulescent giant rosette genus Espeletia in the Andes.We report here the differences in supercooling capacity andcold injury in several Espeletia schultzii populations growingalong an altitudinal gradient. The relationships between supercooling,water potential and leaf anatomy were also investigated. Thesupercooling capacity increased and injury temperature decreasedfrom lower to higher elevation populations. These changes maybe explained in terms of physiological, morphological and anatomicalcharacteristics of the leaves. Key words: Espeletia schultzii, supercooling, freezing avoidance mechanisms     

Characterization and Quantification of Intrinsic Ice Nucleators in Winter Rye (Secale cereale) Leaves

Extracellular ice formation in frost-tolerant organisms is often initiated at specific sites by ice nucleators. In this study, we examined ice nucleation activity (INA) in the frost-tolerant plant winter rye (Secale cereale). Plants were grown at 20[deg]C, at 5[deg]C with a long day, and at 5[deg]C with a short day (5[deg]C-SD). The threshold temperature for INA was -5 to -12[deg]C in winter rye leaves from all three growth treatments. Epiphytic ice nucleation-active bacteria could not account for INA observed in the leaves. Therefore, the INA must have been produced endogenously. Intrinsic rye ice nucleators were quantified and characterized using single mesophyll cell suspensions obtained by pectolytic degradation of the leaves. The most active ice nucleators in mesophyll cell suspensions exhibited a threshold ice nucleation temperature of -7[deg]C and occurred infrequently at the rate of one nucleator per 105 cells. Rye cells were treated with chemicals and enzymes to characterize the ice nucleators, which proved to be complexes of proteins, carbohydrates, and phospholipids, in which both disulfide bonds and free sulfhydryl groups were important for activity. Carbohydrates and phospholipids were important components of ice nucleators derived from 20[deg]C leaves, whereas the protein component was more important in 5[deg]C-SD leaves. This difference in composition or structure of the ice nucleators, combined with a tendency for more frequent INA, suggests that more ice nucleators are produced in 5[deg]C-SD leaves. These additional ice nucleators may be a component of the mechanism for freezing tolerance observed in winter rye.     

Complex bud architecture and cell‐specific chemical patterns enable supercooling of Picea abies bud primordia

Bud primordia of Picea abies, despite a frozen shoot, stay ice free down to ?50 °C by a mechanism termed supercooling whose biophysical and biochemical requirements are poorly understood. Bud architecture was assessed by 3D—reconstruction, supercooling and freezing patterns by infrared video thermography, freeze dehydration and extraorgan freezing by water potential measurements, and cell‐specific chemical patterns by Raman microscopy and mass spectrometry imaging. A bowl‐like ice barrier tissue insulates primordia from entrance by intrinsic ice. Water repellent and densely packed bud scales prevent extrinsic ice penetration. At ?18 °C, break‐down of supercooling was triggered by intrinsic ice nucleators whereas the ice barrier remained active. Temperature‐dependent freeze dehydration (?0.1 MPa K^?1) caused accumulation of extraorgan ice masses that by rupture of the shoot, pith tissue are accommodated in large voids. The barrier tissue has exceptionally pectin‐rich cell walls and intercellular spaces, and the cell lumina were lined or filled with proteins, especially near the primordium. Primordial cells close to the barrier accumulate di, tri and tetrasaccharides. Bud architecture efficiently prevents ice penetration, but ice nucleators become active inside the primordium below a temperature threshold. Biochemical patterns indicate a complex cellular interplay enabling supercooling and the necessity for cell‐specific biochemical analysis.     

Analysis of supercooling activity of tannin-related polyphenols

Based on the discovery of novel supercooling-promoting hydrolyzable gallotannins from deep supercooling xylem parenchyma cells (XPCs) in Katsura tree (see Wang et al. (2012) [38]), supercooling capability of a wide variety of tannin-related polyphenols (TRPs) was examined in order to find more effective supercooling-promoting substances for their applications. The TRPs examined were single compounds including six kinds of hydrolyzable tannins, 11 kinds of catechin derivatives, two kinds of structural analogs of catechin and six kinds of phenolcarboxylic acid derivatives, 11 kinds of polyphenol mixtures and five kinds of crude plant tannin extracts. The effects of these TRPs on freezing were examined by droplet freezing assays using various solutions containing different kinds of identified ice nucleators such as the ice nucleation bacterium (INB) Erwinia ananas, the INB Xanthomonas campestris, silver iodide and phloroglucinol as well as a solution containing only unintentionally included unidentified airborne ice nucleators. Among the 41 kinds of TRPs examined, all of the hydrolyzable tannins, catechin derivatives, polyphenol mixtures and crude plant tannin extracts as well as a few structural analogs of catechin and phenolcarboxylic acid derivatives exhibited supercooling-promoting activity (SCA) with significant differences (p > 0.05) from at least one of the solutions containing different kinds of ice nucleators. It should be noted that there were no TRPs exhibiting ice nucleation-enhancing activity (INA) in all solutions containing identified ice nucleators, whereas there were many TRPs exhibiting INA with significant differences in solutions containing unidentified ice nucleators alone. An emulsion freezing assay confirmed that these TRPs did not essentially affect homogeneous ice nucleation temperatures. It is thought that not only SCA but also INA in the TRPs are produced by interactions with heterogeneous ice nucleators, not by direct interaction with water molecules. In the present study, several TRPs that might be useful for applications due to their high SCA in many solutions were identified.     

Supercooling Capacity Increases from Sea Level to Tree Line in the Hawaiian Tree Species Metrosideros polymorpha

Population-specific differences in the freezing resistance of Metrosideros polymorpha leaves were studied along an elevational gradient from sea level to tree line (located at ca. 2500 m above sea level) on the east flank of the Mauna Loa volcano in Hawaii. In addition, we also studied 8-yr-old saplings grown in a common garden from seeds collected from the same field populations. Leaves of low-elevation field plants exhibited damage at -2 degrees C, before the onset of ice formation, which occurred at -5.7 degrees C. Leaves of high-elevation plants exhibited damage at ca. -8.5 degrees C, concurrent with ice formation in the leaf tissue, which is typical of plants that avoid freezing in their natural environment by supercooling. Nuclear magnetic resonance studies revealed that water molecules of both extra- and intracellular leaf water fractions from high-elevation plants had restricted mobility, which is consistent with their low water content and their high levels of osmotically active solutes. Decreased mobility of water molecules may delay ice nucleation and/or ice growth and may therefore enhance the ability of plant tissues to supercool. Leaf traits that correlated with specific differences in supercooling capacity were in part genetically determined and in part environmentally induced. Evidence indicated that lower apoplastic water content and smaller intercellular spaces were associated with the larger supercooling capacity of the plant's foliage at tree line. The irreversible tissue-damage temperature decreased by ca. 7 degrees C from sea level to tree line in leaves of field populations. However, this decrease appears to be only large enough to allow M. polymorpha trees to avoid leaf tissue damage from freezing up to a level of ca. 2500 m elevation, which is also the current tree line location on the east flank of Mauna Loa. The limited freezing resistance of M. polymorpha leaves may be partially responsible for the occurrence of tree line at a relatively low elevation in Hawaii compared with continental tree lines, which can be up to 1500 m higher. If the elevation of tree line is influenced by the inability of M. polymorpha leaves to supercool to lower subzero temperatures, then it will be the first example that freezing damage resulting from limited supercooling capacity can be a factor in tree line formation.     

Ice nucleation studies of two beetles from sub-antarctic South Georgia

Summary Supercooling points of adults and larvae of the coleopterans Hydromedion sparsutum and Perimylops antarcticus at South Georgia ranged from -3.0 to -5.4°C with Perimylops freezing at c.1.6°C lower than Hydromedion. Intact excised guts from adults of both species froze c. 1°C lower than the adult insects. Ice nucleating activity of homogenized faeces from larvae and adults of both species and excised guts were compared with three potential food plants using an ice nucleation spectrometer. Mean supercooling points of the insect materials at four concentrations in distilled water (range from 0.01 to 10 g 1^–1) were significantly different (P<0.01) within species, and within life stages between species. Differences in the supercooling points of suspensions of Polytrichum alpinum (moss) and Usnea fasciata (lichen) were not significant. In general, differences between supercooling points were greater at the higher concentrations. Histograms of the supercooling points showed unimodal distributions particularly at high concentrations and greater dispersion with increased dilution. Spectra showing the concentration of active ice nucleators over the temperature range 0 to -20°C were developed. These showed that nucleation occurred as high as -2°C in faecal material and all insect samples nucleated above -3°C, whereas the plant materials nucleated between -4 and -5°C. The calculated number of ice nucleators for each material in suspension revealed low values (5.3 to 5.8 × 10³) for the plants, but a greater abundance (1.3 × 10⁵ to 1.3 × 10⁶) in the insect samples. It is concluded that c.1000 active nucleators g^–1 are required for ice nucleation to occur in these suspensions. Ice nucleator activity of a suspension of Hydromedion faeces was much reduced by heating to 75°C, suggesting a proteinaceous structure. These results are discussed in relation to ice nucleation in other insects, and it is concluded that bacteria may be responsible for the high nucleation temperatures, and hence poor supercooling, in these South Georgia insects. An empirical model is developed for ice nucleation spectra based on these data.     

Maintenance of the supercooled state in the gut fluid of overwintering pyrochroid beetle larvae, Dendroides canadensis : role of ice nucleators and antifreeze proteins

  The effect of gut fluid ice nucleators and antifreeze proteins on maintenance of supercooling was explored in fire-colored beetle larvae, Dendroides canadensis, via seasonal monitoring of supercooling points, antifreeze protein activity and ice nucleator activity of gut fluid and/or larvae. During cold hardening in the field, freeze-avoiding larvae evacuated their guts and depressed larval supercooling points. Analysis of gut fluid indicated supercooling points and ice nucleator activity decreased, whereas antifreeze protein activity increased as winter approached. Suspensions of bacteria isolated from guts of feeding larvae collected in spring/summer had higher supercooling points than those from midwinter-collected non-feeding larvae, suggesting bacterial ice nucleators are removed from midwinter gut fluid. The ice nucleation active bacterium Pseudomonas fluorescens was isolated from gut fluid of feeding larvae but was absent in winter. When mixed with purified D.␣canadensis hemolymph antifreeze proteins (structurally similar and/or identical to those in gut fluid), the cumulative ice nucleus spectra of P. fluorescens suspensions were shifted to lower temperatures indicating an inhibitory effect on the bacteria's ice-nucleating phenotype. By extending larval supercooling capacity, both gut clearing and masking of bacterial ice nucleators by antifreeze proteins may contribute to overwintering survival in supercooled insects. Accepted: 8 August 1996     

Change in overwintering mechanism of the Cucujid beetle, Cucujus clavipes

Overwintering larvae of the Cucujid beetle, Cucujus clavipes, were freeze tolerant, able to survive the freezing of their extracellular body fluids, during the winter of 1978–1979. These larvae had high levels of polyols (glycerol and sorbitol), thermal hysteresis proteins and haemolymph ice nucleators that prevented extensive supercooling (the supercooling points of the larvae were ? 10°C), thus preventing lethal intracellular ice formation. In contrast, C. clavipes larvae were freeze suspectible, died if frozen, during the winter of 1982–1983, but supercooled to ～ ? 30°C. The absence of the ice nucleators in the 1982–1983 larvae, obviously essential in the now freeze-susceptible insects, was the major detected difference in the larvae from the 2 years. However, experiments in which the larvae were artifically seeded at ? 10°C (the temperature at which the natural haemolymph ice nucleators produced spontaneous nucleation in the 1978–1979 freeze tolerant larvae) demonstrated that the absence of the ice nucleators was not the critical factor, or at least not the only critical factor, responsible for the loss of freeze tolerance in the 1982–1983 larvae. The lower lethal temperatures for the larvae were approximately the same during the 2 winters in spite of the change in overwintering strategy.     

Zero-sized effect of nano-particles and inverse homogeneous nucleation. Principles of freezing and antifreeze

It was found that freezing of water in terms of homogeneous nucleation of ice never occurs even in ultra-clean micro-sized water droplets under normal conditions. More surprisingly, at sufficiently low supercoolings, foreign nano-particles exert no effect on the nucleation barrier of ice; it is as if they physically "vanished." This effect, called hereafter the "zero-sized" effect of foreign particles (or nucleators), leads to the entry of a so-called inverse homogeneous-like nucleation domain, in which nucleation is effectively suppressed. The freezing temperature of water corresponds to the transition temperature from the inverse homogeneous-like nucleation regime to foreign particle-mediated heterogeneous nucleation. The freezing temperature of water is mainly determined by (i) the surface roughness of nucleators at large supercoolings, (ii) the interaction and structural match between nucleating ice and the substrate, and (iii) the size of the effective surface of nucleators at low supercoolings. Our experiments showed that the temperature of -40 degrees C, commonly regarded as the temperature of homogeneous nucleation-mediated freezing, is actually the transition temperature from the inverse homogeneous-like nucleation regime to foreign particle-mediated heterogeneous nucleation in ultra-clean water. Taking advantage of inverse homogeneous-like nucleation, the interfacial tensions between water and ice in very pure water and antifreeze aqueous solutions were measured at a very high precision for the first time. The principles of freezing promotion and antifreeze and the selection for the biological ice nucleation and antifreeze proteins are obtained. The results provide completely new insights into freezing and antifreeze phenomena and bear generic implications for all crystallization systems.     

Nitrate Supply and the Biophysics of Leaf Growth in Salix viminalis

The influence of nitrogen on leaf area development and the biophysicsof leaf growth was studied using clonal plants of the shrubwillow, Salix viminalis grown with either optimal (High N) orsub-optimal (Low N) supplies of nitrate. Leaf growth rate andfinal leaf size were reduced in the sub-optimal treatment andthe data suggest that in young rapidly growing leaves, thiswas primarily due to changes in cell wall properties, sincecell wall extensibility (% plasticity) was reduced in the LowN plants. The biophysical regulation of leaf cell expansion also differedwith nitrogen treatment as leaves aged. In the High N leaves,leaf cell turgor pressure (P) increased with age whilst in theLow N leaves P declined with age, again suggesting that foryoung leaves, cell wall plasticity limited expansion in theLow N plants. Measurements of cell wall properties showed thatcell wall elasticity (%E) was not influenced by nitrogen treatmentand remained constant regardless of leaf age. Key words: Salix, cell wall extensibility, nitrogen nutrition, biophysics of leaf growth     

Freezing Characteristics of Rigid Plant Tissues (Development of Cell Tension during Extracellular Freezing)

The freezing characteristics and development of cell tension during extracellular freezing were examined in supercooling stem tissues of riverbank grapes (Vitis riparia) and cold-hardened leaves of live oak (Quercus virginiana) and mountain cranberry (Vaccinium vitis-idaea). Dormant stem xylem and pith tissues of river-bank grapes were resistant to freeze-induced dehydration above the homogeneous nucleation temperature, and they developed cell tension reaching a maximum of 27 MPa. Similarly, extracellular freezing induced cell tension in the leaves of live oak and mountain cranberry. Maximum cell tension in the leaves of live oak was 16.8 MPa and 8.3 MPa in the leaves of mountain cranberry. Following peak tensions in the leaves, a decline in the pressure was observed with progressive freezing. The results suggest that resistance to cell deformation during extracellular freezing due to cell-wall rigidity can lead to reduced cell dehydration and increased cell tension. A relationship to predict freezing behavior in plant tissues based on cell rigidity is presented. Based on cell-water relations and ice nucleation rates, cell-wall rigidity has been shown to effect the freezing characteristics of plant tissues, including freeze-induced dehydration, supercooling, and homogeneous nucleation temperatures.     

Freezing avoidance in Andean giant rosette plants

Abstract Frost avoidance mechanisms were studied in Espeletia spicata and Espeletia timotensis, two Andean giant rosette species. The daily courses of soil, air and tissue temperatures were measured at a site at circa 4000 m. Only the leaves were exposed to subzero temperatures; the apical bud and stem pith tissues were insulated by surrounding tissues. The leaf tissues avoided freezing by supercooling rather than by undergoing active osmotic changes. The temperatures at which ice formed in the tissues (the supercooling points) coincided with injury temperatures indicating that Espeletia tissue does not tolerate any kind of ice formation. For insulated tissue (apical bud, stem pith, roots) the supercooling point was around - 5°C coinciding with the injury temperature. Supercooling points of about –13 to - 16°C were observed for leaves. These results contrast with those reported for Afroalpine giant rosettes which tolerate extracellular freezing. The significance of different adaptive responses of giant rosettes to similar cold tropical environments is discussed.     

Freezing of Plant Tissues3

The freezing of plant cells under the microscope was studiedon three types of tissues: Tradescantia staminal hairs, fruitskin, and moss leaf. Cinemicrography was used to record stagesin the freezing processes, which are otherwise un observable.The following phenomena have been demonstrated and discussed:(1) The protective effect of persistent supercooling. (2) Ice-inoculationof cells and factors affecting it. (3) Sequence of ice formationwithin cells. (4) Modes of ice formation as affected by supercooling.(5) Freezing of cell walls and their longitudinal splitting.     

Bulbing in Onions: Photoperiod and Temperature Requirements and Prediction of Bulb Size and Maturity

Bulb size and maturity are key characteristics of an onion cropand the onset of bulbing is an important determinant of these.In this paper we describe an experiment in which bulb and neckdiameter and leaf number were measured in onion crops (cultivarsPukekohe Longkeeper and Early Longkeeper) with different sowingdates planted at two different locations in New Zealand. A sensitiveindicator of earliest time of bulbing was developed using theratio of bulb and neck diameters and the statistical techniqueof cusums. Bulb diameter at bulbing was related to thermal timeaccumulated prior to bulbing. Bulbing only occurred when dualthresholds of a minimum thermal time of 600 degree days anda photoperiod of 13.75 h were reached. Mathematical relationshipswere developed between leaf number, sowing date, bulbing dateand bulb growth and maturity. Final bulb size could be predictedfrom bulb size at bulbing and number of leaves produced afterbulbing. Bulb maturity date could be predicted by number ofleaves after bulbing. Allium cepa L.; onion; temperature; photoperiod; bulb-neck ratio; leaf number; bulbing