Short-term responses of Brassica rapa plants to low root temperature: Effects on nitrate uptake and its translocation to the shoot

Brassica rapa L. plants were grown hydroponically for 5 or 6 weeks at 20°C and then half batches of plants were transferred to tanks in which the root temperature was lowered decrementally over 1 h to 7°C. Changes in nitrate uptake rate (NUR) and nitrate transfer from roots were studied in relation to transpiration and root pressure xylem exudation flow rates over a 48- or 72-h period. The response of plants following the root temperature decrease was biphasic. During phase 1, NUR and water and solute flow rates through the root decreased sharply. Coping mechanisms came into operation during phase 2, and tended to offset the effects of low temperature. The 3-h cold-treated roots exhibited a very low NUR but 48-h cold-treated roots partly recovered their ability to absorb nitrate. Transpiration rate decreased more slowly (during 24 h) than both root xylem exudation and parameters of root conductivity (during 6 h). Beyond these respective times, transpiration rate was balanced while root xylem exudation clearly increased, but without returning to the level of control plants. Nitrate transfer to the root xylem was strongly and rapidly affected by low root temperature, but the subsequent readjustment was such that no or little difference compared with the control was apparent after 48 h. Water and solute flows were strongly decreased when nitrate was replaced by chloride in the culture solution during exudation sampling. The major role of nitrate in root hydraulic conductivity and root xylem exudation is discussed.     

Low-temperature pretreatment of the root system of Brassica rapa L. plants: effects on the xylem sap exudation and on the nitrate absorption rate

Brassica rapa plants were exposed for a 52 h period (as pretreatment) to a differential temperature (DT) between roots (5°C) and shoots (20°C), while control plants were maintained with both shoot and roots at 20°C (warm grown = WG). Measured at 20°C, volume flow of xylem exudate from roots of DT plants was enhanced compared with that from WG plants, while transpiration flows were similar in pretreated and control plants. Both transpiration and exudation flows were dependent upon shoot/root ratio. Differences in the volume flow of exudate were principally related to increases in root hydraulic conductance. Anion fluxes (notably nitrate) into xylem exudate of DT plants were significantly greater than those into exudate of WG plants. This enhancement of nitrate flow from the pretreated roots was associated with a two-fold increase in nitrate uptake rate. The relationship of the cold-induced change in nitrate uptake capacity with shoot/root ratio is discussed in terms of control of nitrate absorption by shoot sink strength.     

Biomass, reproductive output, and physiological responses of rapid-cycling Brassica (Brassica rapa) to ozone and modified root temperature

Brassica rapa L. (rapid-cycling Brassica), was grown in environmentally controlled chambers to determine the interactive effects of ozone (O₃) and increased root temperature (RT) on biomass, reproductive output, and photosynthesis. Plants were grown with or without an average treatment of 63 ppb O₃. RT treatments were 13°C (LRT) and 18°C (HRT). Air temperatures were 25°C/15°C day/night for all RT treatments.
Ozone affected plant biomass more than did root temperature. Plants in O₃ had significantly smaller total plant d. wt, shoot weight, leaf weight, leaf area and leaf number than plants grown without O₃. LRT plants tended to have slightly smaller total plant d. wt, shoot weight, root weight, leaf weight, leaf area, and leaf number than HRT plants. For all variables, LRT plants grown in O₃ had the smallest biomass, and plants grown in HRT without O₃ had the largest biomass.
Ozone reduced both fruit weight and fruit number; LRT also reduced fruit weight but had no effect on fruit number. Ozone reduced photosynthesis but RT had no effect. Conductance and internal CO₂ were unaffected by O₃ or RT.
These studies indicate that plant growth with LRT might be more reduced in the presence of O₃ than growth in plants with HRT, which might be able to compensate for O₃-caused reductions in photosynthesis to avoid decreased biomass and reproductive output.     

脱落酸对低温下雷公藤幼苗光合作用及叶绿素荧光的影响

以1年生雷公藤扦插苗为试材,研究低温胁迫下不同浓度外源脱落酸（ABA,0、5、10、15、20、25 mg·L^-1)叶面喷施处理对雷公藤叶片光合作用及叶绿素荧光参数的影响.结果表明：喷施20 mg·L^-1的ABA能显著提高雷公藤幼苗的抗冷性,减缓低温下雷公藤叶片净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(g_s)、胞间CO₂浓度(C_i)的下降幅度,提高幼苗叶片的光合能力.低温处理6 d后,随着ABA浓度上升,雷公藤叶片的初始荧光(F_o)下降,最大荧光(F_m)和PSII最大光化学效率(F_v/F_m)上升,PSII实际光化学量子产量(Φ_PSⅡ)、光化学猝灭系数(q_P)先下降后上升,而非光化学猝灭系数(q_N)呈下降－上升－下降趋势.P_n、g_s、q_P、F_m和F_v/F_m均在20 mg·L^-1ABA处理时达到峰值.不同浓度ABA的相对电子传递速率(rETR)随着光化光强度增加呈先上升后下降的趋势,当光化光强度(PAR)达到395 μmol·m^-2s^-1时,各处理的rETR达到最高值,其中25 mg·L^-1和20 mg·L^-1ABA处理分别比对照高17.1%和5.2%.雷公藤叶片Φ_PSⅡ的光响应曲线均随光化光强度升高而下降,q_N的光响应曲线则呈相反趋势.     

Plasmodiophora brassicae-induced Cell Death and Medium Alkalization in Clubroot-resistant Cultured Roots of Brassica rapa

Plasmodiophora brassicae causes clubroot in the turnip, Brassica rapa L. We used organ cultures of adventitious roots from B. rapa seedlings to investigate the initial response of resistant and susceptible cultivars to P. brassicae infection. Primary plasmodia of P. brassicae were observed in root hairs of both susceptible and resistant cultured roots. On the other hand, secondary plasmodia were able to proliferate only in the susceptible root culture but not in the resistant one. Root cultures from the susceptible cultivar all developed clubroot 4 weeks after treatment with 10⁴, 10⁵ or 10⁶ spores/ml, but roots from the resistant cultivar did not develop clubroot under the same conditions. Cell death, as measured by Evans blue and TTC dye methods, was observed in cultured roots from the resistant cultivar but did not occur in roots from the susceptible cultivar after exposure to P. brassicae spores. Cell death was inhibited almost completely by EGTA and verapamil but not by the calmodulin antagonist W7. These results suggest the involvement of Ca²⁺ in P. brassicae‐induced cell death. Alkalization of the root culture medium of the resistant cultivar was observed 2 days after treatment with P. brassicae spores but was not observed in root culture medium from the susceptible strain. We conclude that our root culture system must be a useful tool for further studies of the molecular mechanism of clubroot resistance.     

低温胁迫下脱落酸对线粒体膜磷脂酶D活性的影响

高山离子芥(Choraspora bungeana)是一种稀有高山冰缘植物,其生活环境具有低温、强紫外线等胁迫因子。PLD在膜磷脂降解及磷脂信号转导过程中发挥着重要作用,但其活性往往受到多种因素的影响。该研究以高山离子芥试管苗为材料,研究了4℃、0℃和-4℃胁迫下,ABA对高山离子芥试管苗叶中线粒体膜结合态PLD活性的影响。结果表明:10,50和100μmol·L~(-1)脱落酸(ABA)处理高山离子芥后,线粒体膜结合态PLD活性均较未添加ABA的处理组线粒体膜结合态PLD活性高,其中以50μmol·L~(-1) ABA对离子芥叶中线粒体膜结合态PLD活性的促进作用最为显著;外施0.3 mmol·L~(-1)的ABA合成抑制剂钨酸钠处理高山离子芥后,线粒体膜结合态PLD活性较对照组线粒体膜结合态PLD活性降低;在50μmol·L~(-1) ABA+5 mmol·L~(-1) EGTA处理组中,高山离子芥叶中线粒体膜结合态PLD活性低于未添加EGTA处理组线粒体膜结合态PLD活性;在0.3 mmol·L~(-1)钨酸钠+10 mmol·L~(-1)CaCl_2处理组中,高山离子芥叶中线粒体膜结合态PLD活性高于未添加CaCl_2处理组线粒体膜结合态PLD活性。由此推测,低温胁迫下ABA可能通过Ca~(2+)介导影响高山离子芥叶中线粒体膜结合态PLD的活性。     

Effects of abscisic acid,low temperature,and plant age on cytoskeleton and phosphorylated proteins

The effects of exogenous abscisic acid (ABA), low temperature, and seedling age on the content of tubulin, actin, and phosphorylated proteins and the structural organization of microtubules (MTs) in cells of different tissues and organs of winter wheat cultivars contrasting in cold hardiness were studied by immunocytochemical methods using monoclonal (against - and -tubulin and actin) and polyclonal (phosphothreonine) antibodies. The leaves and roots of five- and nine- day-old seedlings of three cultivars were characterized by unequal proportion of actin/tubulin proteins. ABA decreased the content of the cytoskeleton and the 60-kD phosphorylated proteins, thus promoting a decrease in the number of MTs and occurrence of a less branched network of weakly fluorescent tubulin components in the cells of the root differentiating zone (which is most responsible for the development of cold hardiness in wheat). Although the cold acclimation of plants (3°C, 7 days) did not change the level of tubulin and actin proteins, it evoked the spatial aggregation of MT, leading to formation of a dense network of tubulin cytoskeleton comprised of thick bundles of intensively fluorescent MTs. In the case of a combined action of the studied factors, low temperatures abolished the hormone effect described above, evoking an increase in the content of the cytoskeletal and 60-kD phosphorylated proteins and MT structures. We suggest that the ABA-induced decrease in the levels of proteins and MTs occurs at the initial stages of plant cold acclimation (3°C, 2-3 days). It may be the signal that triggers the processes of low-temperature adaptation. As the duration of cold acclimation increased (3°C, 7 days), the role of ABA in the formation of plant tolerance decreased. Apparently, in this case other hormone-independent mechanisms of frost hardiness development are triggered, in which the role of the cytoskeleton components and cytoskeleton-associated proteins increases.     

温度对大菜粉蝶生长发育的影响

为明确温度对大菜粉蝶生长发育的影响,在16、20、22、25、28和31℃共6个恒温条件下以白菜为寄主进行饲养,测定了温度对大菜粉蝶各虫态的发育历期、发育速率、存活率及成虫寿命等的影响,并采用直接最优法计算其各虫态的发育起点温度和有效积温。结果表明：在16～28℃温度范围内,大菜粉蝶的卵、幼虫、蛹及全世代平均发育速率具有随温度升高而加快的趋势,存活率则随着温度的升高而降低;卵、幼虫、蛹和成虫的发育起点温度分别为11.76℃、6.87℃、5.30℃和14.91℃,有效积温分别为38.14、213.90、169.68和43.01 d·℃。在28℃时大菜粉蝶卵的孵化率、各龄幼虫存活率、化蛹率、蛹重及羽化率均显著低于其他温度,31℃条件下不能完成个体发育。由此可见,高温环境不利于大菜粉蝶的存活和生长发育。研究结果可为大菜粉蝶田间发生期预测及综合防治提供科学依据。     

The effect of abscisic acid on root and shoot growth of cauliflower plants

Addition of abscisic acid (ABA) to the nutrient solution increased the root to shoot ratio of hydroponically-grown cauliflower plants by reducing the dry weight of the shoot and increasing that of the root. At concentrations higher than 10^–7 M, ABA increased root branching and root hair formation. Root extension was inhibited in plants kept continuously in solutions containing high ABA concentrations but following removal from the ABA solution root elongation was increased in comparison with plants given no ABA treatment. This elongation was greatest in plants with increased root branching caused by higher ABA concentrations.     

Fatty acid desaturation in monogalactosyldiacylglycerol of Brassica napus leaves during low temperature acclimation

Brassica napus plants grown at 5°C have the potential to desaturate fatty acids in the major membrane diacylglycerols of leaves at rates much higher than those of plants grown at 30°C. This low temperature-induced desaturation (LTD) is rapidly deactivated if plants grown at 5°C are transferred to 30°C for several hours. By exposure to ¹⁴CO₂ and a computer simulation program, we estimated the rate of desaturation of monogalactosyldiacylglycerol by (ω9-, ω6- and ω3-desaturases of plants grown at 5, 10, 20 and 30°C. Data show that LTD can be induced in mature leaves of plants grown at 20 and 30°C after transfer to 5°C. Full activation of LTD takes several weeks and occurs more rapidly in plants grown at 20°C than 30°C. This activation is largely due to the increased activity of ω9- and ω6-desaturases on C16 fatty acids and ω6-desaturase on C18 fatty acids.     

Effects of short chain fatty acids on radicle emergence and root growth in lettuce

Abstract. Short chain fatty acids inhibit both radicle emergence and root growth in lettuce. The transition from ineffectual to inhibitory levels occurs abruptly. Root growth is more sensitive to lower concentrations than radicle emergence and is invariant with chain length. The effect of short chain alcohols on radicle emergence is similar to that of short chain acids, but their comparatively severe inhibition of root growth varies with chain length. Alkanes of the same chain lengths have no noticeable effect. Respiration is not altered by a representative short chain fatty acid (heptanoic). Lettuce seeds are sensitized to phytochrome-absorbed light by short chain fatty acids as found by Berrie and co-workers.     

与大白菜霜霉病抗性主效QTL连锁的分子标记开发

霜霉病是危害大白菜的三大病害之一,该病的发生会严重影响大白菜的产量及品质,因而研究与霜霉病抗性QTL紧密连锁的分子标记对大白菜抗病新品种培育具有重要意义。该研究在前期工作的基础上,选用高感霜霉病株系91-112、高抗霜霉病株系T12-19以及由二者为双亲构建的DH群体为实验材料,针对大白菜霜霉病抗性主效QTL——BrDW所在的标记区间,利用已有的大白菜基因组信息发展与抗性QTL紧密连锁的分子标记,通过Blast和IMap分析,将与BrDW连锁的RAPD标记K14-1030定位于大白菜KBrB058M10上(位于Contig214上),根据KBrB058M10附近的BAC及BAC-end序列设计引物,结合限制性内切酶酶切及HRM分析方法,筛选得到5个与BrDW连锁的分子标记,包括1个Indel标记Brb062-Indel230,3个CAPS标记Brb094-DraⅠ787、Brb094-AatⅡ666和Brb043-BglⅡ715,1个SNP标记Brh019-SNP137;同时,通过筛选与目标区域具有同源性的Unigene序列得到了1个与BrDW紧密连锁的SSR标记bru1209。标记Brb062-Indel230、Brb094-DraⅠ787、Brb094-AatⅡ666、Brb043-BglⅡ715、Brh019-SNP137和bru1209与RAPD标记K14-1030之间的遗传距离分别为4.3 cM、1.7 cM、5.9 cM、5.9 cM、4.6 cM和0.8 cM,在对DH群体中的抗性株系选择上准确率分别为69.7%、70.9%、72.4%、72.4%、58.3%和74.2%,可应用于分子标记辅助选择,为霜霉病抗性分子育种奠定了良好基础。     

‘津田’芜菁Transparent Testa Glabra 1（BrTTG1）基因的克隆及表达分析

TTG1（Transparent Testa Glabra 1）蛋白是一种WD40类蛋白,参与植物的生长和发育。采用RT-PCR方法从芜菁品种‘津田＇中克隆了BrTTG1 cDNA序列（GenBank登录号HM208590）。该基因cDNA开放阅读框长度为1 014 bp,编码一个由337个氨基酸残基组成的蛋白,该蛋白分子量为37.28 kDa,理论等电点为4.66。与其他植物中的TTG1蛋白进行同源性比对结果显示,BrTTG1与甘蓝型油菜的TTG1同源性最高。BrTTG1蛋白在31～337位氨基酸处含有WD40超家族的保守结构域。荧光定量PCR检测BrTTG1在‘津田＇芜菁不同组织中的表达结果表明,该基因在有花青素合成的红色‘津田＇芜菁根皮中表达量最高。     

Commercially available plant growth regulators and promoters modify bulk tissue abscisic acid concentrations in spring barley, but not root growth and yield response to drought

Field and lysimeter experiments were conducted in 2002 to investigate the effects of an antigibberellin growth regulator (Moddus, active ingredient trinexapac‐ethyl, Syngenta Crop Protection UK Ltd, Whittlesford, Cambridge, UK) and an auxin‐stimulating (Route, active ingredient zinc ammonium acetate, De Sangosse Ltd, Swaffham Bulbeck, Cambridge, UK) growth promoter on root growth, soil water extraction and the drought response of spring barley. The effects on root growth and distribution were investigated in the field. The effects on the drought response were studied in 1.2‐m‐deep lysimeters packed with a loamy sand subsoil and sandy loam topsoil. Lysimeters were located under a fixed rain shelter, and drought was imposed by withholding irrigation. In both field and lysimeter experiments, growth regulator/promoters were applied to cv. Optic at early tillering according to the manufacturers’ recommendations. After withholding irrigation from lysimeters at Zadoks growth stage (GS) 21 (37 days after sowing), 50% of the profile available water had been depleted by flag leaf emergence (GS 37/39; 62 days after sowing). Drought significantly reduced stem biomass at ear emergence (GS 59; 78 days after sowing) but not leaf or ear dry weight; this was before there was any significant reduction in leaf water potential or stomatal conductance to water vapour. The reduction in stem biomass may reflect a change in partitioning between shoot and root in response to soil drying. When averaged over growth regulator/promoter treatments, drought reduced grain yield by approximately 1 t ha^?1. This was associated with a reduction in both ears per m² and grains per ear. The mean grain weight was not reduced by drought, in spite of significant reductions in stomatal conductance and canopy lifespan post‐anthesis. Route, and to a lesser extent Moddus, significantly increased abscisic acid accumulation in the stem base of droughted plants, and there was some indication of a possible delay in stomatal closure in Route‐treated plants as the soil moisture deficit developed. However, there was no significant effect on the amount of soil water extracted or grain yield under drought. Similarly, in field experiments, neither Route nor Moddus significantly altered total root length, biomass or distribution. There is little evidence from these experiments or in the literature to support the use of antigibberellin or auxin‐simulating growth regulator/promoters to modify root growth and drought avoidance of spring barley.     

外源ABA对低温胁迫水稻过氧化物酶同工酶的影响

采用聚丙烯酰胺垂直板电泳法,就外源ABA对低温胁迫下水稻幼苗过氧化物酶（POD）同工酶的影响进行了研究。结果表明,低温胁迫下,水稻幼苗长势明显下降,与其相应的过氧化物酶同工酶也发生了变化,外施ABA(10^-5mol/L）提高了水稻幼苗的抗寒能力,其相应的过氧化物酶同工酶却表现为部分酶活性增强,说明：（1）过氧化物同工酶的变化将引起植物对低温的抗性反应;（2）外源ABA引起自由基清除酶（如POD）活力的升高,是植物抗寒性增强的原因之一。     

白菜和甘蓝基因组转座子表达及其对基因调控的潜在影响

转座子或转座元件是大多数真核生物基因组的主要组成成分。甘蓝(Brassica oleracea)基因组比白菜(B. rapa)大主要是转座子的扩增差异造成的。然而, 这两个芸薹属近缘物种转座子表达水平以及对基因的调控和功能的影响目前还不清楚。文章对白菜和甘蓝叶、根、茎3个器官的转录组数据进行了初步分析。结果显示, 转座子的表达量很低, 转录组reads中有1%来自转座子的转录本; 转座子的表达存在器官差异, 且不同类别和家族的转座子表达量相差很大, 相同类别和同一家族的转座子在白菜和甘蓝基因组中的表达活性也不相同。进一步鉴定到转录读出的LTR反转座子, 其与下游基因距离小于2 kb的有41个, 小于100 bp的有9个, 这些LTR的转录读出很可能通过正义或反义的转录本激活或干扰下游基因的表达。同时, 具有转录读出的intact LTR比solo LTR具有更强的读出活性。通过深入分析转座子的插入位点发现, 白菜基因组中转座子插入基因内部的频率比甘蓝基因组中的高; 与反转座子相比, DNA转座子更偏向于插入或保留在基因的内含子当中。这些结果为认识转座子对其他蛋白编码基因的影响提供了基础。     

低温胁迫下内源脱落酸对马尾松优良种源抗寒性的调节作用

以2个马尾松优良种源苗期的叶片为材料,对系列低温梯度条件下不同种源叶片的内源ABA、内源保护酶系统的动态变化的规律以及相互作用进行研究.结果表明:在降温过程中通过内源ABA的变化来调控内源保护酶的变化趋势,并协调多种酶的相互作用来抵抗低温.内源ABA、SOD对零上低温敏感,属冷害敏感型调节物质,它们在零上低温前达到最高...     

Influence of root-zone temperature on growth,development and yield of cucumber plants cv. Toska

Summary In a first experiment, cucumber transplants (cucumis sativa L. cv. Toska) were grwon at five root-zone temperatures (RZT) ranging from 12° to 36°C. Maximum shoot growth and total leaf area were obtained at 24° and 30°C (RZT). In a second experiment, cucumber transplants were submitted to five RZT (12, 18, 24, 30 and 36°C) and five night air temperatures (NAT) that were maintained either constant at 9°, 13° and 17°C or splitted (in two halfs) at midnight (17°/12°C, 17°/9°C). Root-zone warming to 24° or 30°C increased cucumber plant growth and leaf development, but did not compensate completely the loss of productivity induced by low NAT. Split-night temperature had greater effects under the lowest NAT (17°/9°C) and at high RZT (24° or 30°C). In a third experiment, soil warming caused large increase in yields when cucumber plants were grown in the spring, but had very little effects in the fall.     

Canola seed germination and seedling emergence from pre-hydrated and re-dried seeds subjected to salt and water stresses at low temperatures

Low soil temperatures and low water potentials reduce and delay the seed germination of canola (Brassica rapa L., B. napus L.) in western Canada. Germination is also very sensitive to the salinity effects of nitrogen fertiliser placed with the seed, especially when the seed bed is relatively dry. The effects of pre-hydration and re-drying treatment on canola (Brassica rapa L. cv. Tobin) seed germination and seedling emergence at 10°C subjected to either a water or salt stress were determined. Low water potentials, induced by polyethylene glycol (PEG 8000), low soil moisture, or high concentrations of salts, reduced both germination and seedling emergence, and increased the time to 50% germination and emergence of seeds at 10°C. At equal osmotic potentials, Na₂SO₄ was less inhibitory on low temperature germination than either NaCl or PEG, suggesting that the sulphate ion partially alleviated the inhibitory effects of low water potential. Solutions of NaCI produced more abnormal seedlings compared to Na₂SO₄, suggesting that NaCl was more toxic than Na₂SO₄ during seedling development. Pre-hydration and re-drying partially overcame the inhibitory effects of both low water potential and salts on seed germination and seedling emergence at 10°C. The seed treatment increased the germination rate in Petri dishes and seedling emergence from a sandy loam soil. Water potentials or soil water contents required to inhibit 50% germination or emergence at 10°C were lower for treated seeds compared to control seeds. Salt concentrations inhibiting 50% emergence were higher for treated seeds than control seeds. Neither treated nor control seeds produced seedlings which emerged if the soil water content was lower than 9% or when the soil was continuously irrigated with salt solutions of 100 mmol kg^-1 of NaCl or 50 mmol kg^-1 of Na₂SO₄. These results suggest that the pre-hydration and re-drying treatment did not lower the base water potentials at which seedling emergence could occur. Abnormal seedlings were observed in both treated and control seeds, particularly if the soil was watered with NaCl solutions; however, the seed treatment reduced the number of abnormal seedlings.     

低温胁迫对麻竹叶片和根系抗性生理指标的影响

采用室内人工低温处理,研究了麻竹(Dendrocalamus latiflorus)叶片及根部质膜透性、丙二醛、可溶性蛋白、可溶性糖含量,超氧化物歧化酶(SOD)、过氧化物酶(POD)活性和膜脂脂肪酸组成的变化,寻找与竹类植物耐寒性关系最密切的抗性生理指标。结果表明:低温预处理(8℃)15d后,麻竹叶片中可溶性糖、可溶性蛋白含量、POD活性显著提高,根部可溶性糖含量、POD活性显著升高;低温胁迫处理(-2℃)72h后,经低温预处理的麻竹叶片可溶性糖、可溶性蛋白质含量、SOD、POD活性显著高于未经低温预处理,而质膜透性显著低于未经低温预处理,但膜脂过氧化程度显著高于-2℃处理前;经低温预处理的麻竹根部SOD、POD活性及膜脂不饱和脂肪酸相对含量显著高于未经低温预处理,而质膜透性、膜脂过氧化程度较-2℃处理前无显著差异。说明剧烈降温对麻竹生理特征造成严重影响,叶片通过提高可溶性蛋白、可溶性糖含量,维持较高的POD活性以减轻低温伤害;根系则通过维持较高的SOD、POD活性以减轻低温下膜脂过氧化水平,并通过提高膜脂不饱和脂肪酸比例降低质膜透性来抵御低温对膜的伤害。