不同耐盐性花生品种对NaCl胁迫的光合和抗逆生理响应特征

以较耐盐花生品种‘花育25’、‘鲁花12’和盐敏感品种‘海花1’、‘花育20’为材料,采用盆栽试验,设置0、1.0、2.0、3.0 g/kg土壤NaCl胁迫浓度梯度,测定其净光合速率、表观量子效率、气孔导度等光合特性,以及抗氧化酶活性和渗透调节物质含量等指标,明确NaCl胁迫条件下不同耐盐性花生品种光合和生理生化特性的适应特征。结果表明：(1)NaCl胁迫明显抑制各品种花生叶片光合作用,净光合速率随盐胁迫浓度的升高呈明显降低的趋势。(2)各品种花生叶片净光合速率均先随光照强度的增强而升高,当光强达到一定数值时趋于平稳;光补偿点和光饱点因品种和盐胁迫浓度差异较大,较高的盐胁迫浓度使叶片光补偿点升高,盐敏感品种的光饱和点降低。(3)盐胁迫条件下,各品种叶片表观量子效率和最大净光合速率均随盐胁迫强度的增加呈显著降低趋势,盐敏感品种利用弱光的能力在低盐胁迫下强于耐盐品种,其最大净光合速率在较高盐胁迫浓度(3.0 g/kg)下明显低于耐盐品种,但两类品种的叶片表观量子效率降幅相近(78.65%~88.00%)。(4)在NaCl胁迫下,耐盐品种叶片自由水含量显著高于盐敏感品种;在2.0～3.0 g/kg NaCl胁迫下,耐盐品种叶片SOD、CAT、POD活性和MDA含量的升降幅度均低于盐敏感品种;耐盐品种在NaCl浓度低于2.0 g/kg时的抗氧化能力明显高于盐敏感品种。研究发现,盐胁迫下花生品种抗盐耐逆的主要生理响应特征是提高光补偿点和最大净光合速率,增强叶片持水能力和物质代谢能力,以及提升抗氧化和渗透调节能力。     

大豆种质资源萌发期耐旱性评价

随着全球气候的变化,干旱已成为限制大豆产量的重要环境因素之一,大豆种质资源耐旱性鉴定对大豆耐旱品种的培育及大豆耐旱机理的研究具有重要的意义。本研究首先选择4份耐旱性不同的材料,以PEG-6000作为渗透胁迫剂,研究了大豆种质在0%、10%、15%、16%、18%、20%、30%的PEG-6000溶液的胁迫下,相对发芽率、相对发芽势、相对胚根长度、萌发耐旱指数的变化。结果表明15%~18%PEG-6000溶液是大豆萌发期耐旱鉴定的适宜浓度范围。本研究采用16%PEG-6000溶液对568份大豆资源进行大豆种子萌发期耐旱鉴定,以相对发芽势、相对发芽率、相对胚根长度、相对苗高、萌发耐旱指数、活力指数作为评价指标,应用平均隶属函数评价法对大豆种质萌发期耐旱性进行综合评价,筛选出耐旱种质4份,较耐旱种质18份,中间型种质110份,较敏感种质194份,敏感型种质242份。     

Identification of rice varieties with high tolerance or sensitivity to lead and characterization of the mechanism of tolerance

Pb inhibits plant growth. To study Pb tolerance in rice (Oryza sativa), we screened 229 varieties for Pb tolerance or sensitivity. Three-day-old seedlings were treated for 12 d with 20 microM Pb solution. Based on the dry weight of the root, three Pb-tolerant (var CH-55, var KH-2J, var Kumnung) and three Pb-sensitive (var Aixueru, var C-9491, var Milyang23) rice varieties were selected. The root biomasses of the tolerant varieties were approximately 10-fold higher than those of the sensitive ones. The greatest morphological difference between the two groups was in the growth of the adventitious roots, as tolerant lines were able to develop adventitious roots after 6 d of Pb treatment, whereas sensitive ones did not develop any even after 15 d. The growth of adventitious roots in the tolerant varieties was dependent on a mechanism, whereby Pb was altered to a form that cannot be taken up by the tissue, because (a) the solution in which the tolerant varieties of rice had grown still contained Pb but nevertheless did not affect the root growth of new rice seedlings, and (b) the adventitious roots of tolerant seedlings developed in Pb solution contained little Pb. The oxalate content in the root and root exudate increased upon Pb treatment in the tolerant varieties, whereas the opposite was observed for the sensitive ones. Oxalate added to the growth solution ameliorated the inhibition of root growth by Pb. These results suggest that compounds such as oxalate secreted from the root may reduce the bio-availability of Pb, and that this may constitute an important Pb tolerance mechanism in the tolerant rice varieties studied here.     

水稻资源芽期和苗期耐盐碱性综合评价及耐盐基因分析

培育耐盐碱水稻品种是应对全球人口日益增长的重要途径之一。文中以21份耐盐碱性不同的水稻品种(系)为材料,在芽期和苗期设置6个不同盐碱浓度处理,测定了发芽势、发芽率、芽长、根长、根数、芽鲜重和苗鲜总重等指标,以各指标盐害率的平均值作为耐盐碱性的综合评价标准。结果表明随着盐碱浓度的提升,对种子萌发和生长的抑制越明显。在1%NaCl加0.25%NaHCO3溶液处理下,发芽率盐害率变异最大,为0%–89.80%。在所有浓度处理下,各性状指标的盐害率都具有相似的变化趋势。筛选到4份综合耐盐碱能力强(大酒谷、日本晴、魔王谷和02428)和7份弱的种质资源。比较了4份耐盐碱强和3份耐盐碱弱的资源耐盐基因序列:OsHAL3和OsRR22基因在7份材料中没有差异,SKC1和DST基因在耐盐碱强和耐盐碱弱的品种之间有明显的变异。研究结果为进一步挖掘水稻耐盐碱基因和培育耐盐碱水稻新品种提供了种质资源和理论基础。     

The effects of lime and phosphorus on the function of wheat roots in acid top soils and subsoils

Two wheat varieties with differing aluminium tolerance were grown in pots of acid soil. Liming did not change significantly the amounts of chemically extractable P and K, but caused improved vegetative growth, increased inflow of P and K and reduced uptake of Al. Without lime, roots had a higher content and concentration of P than shoots; liming reversed this. Without lime the sensitive variety with a shorter root length had an Al inflow ten times that of the tolerant one: tolerance involves a mechanism for exlcuding Al. The inflow of P per unit inflow of Al (mol ratio) without lime was three times greater for the tolerant variety which therefore has more P to counteract the effects of Al. The same varieties were grown in two-layer soil columns, with a low P status and a limed topsoil and acid subsoil. Liming the subsoil improved plant growth but this was still restricted by low P availability. Addition of P to the topsoil caused good growth regardless of subsoil acidity: root growth increased in both layers and P (labelled with³²P) taken up from the topsoil was translocated to roots in the subsoil. This P inactivated root Al and allowed the roots to grow and take up more P from the acid subsoil with however a reduction in inflow. The sensitive variety was affected more by the acid subsoil and low P availability, had a similar ability to translocate P to subsoil roots but could not attain the growth rate of the tolerant wheat even with P and lime.     

花生品种耐盐性指标筛选与综合评价

采用盆栽试验,设置不同盐胁迫浓度,通过萌发至幼苗期的出苗速度、植株形态和生物量等指标对200个花生品种(系)进行耐盐性评价.结果表明: 随盐胁迫浓度的增加,花生出苗时间延长,对植株形态建成抑制加重,物质积累减少.鉴定花生品种耐盐性强弱的适宜盐胁迫浓度为0.30%~0.45%.采用隶属函数值法将10个指标归结为平均隶属函数值,根据不同胁迫浓度下各指标与平均隶属函数值之间的相关性大小,植株鲜质量、地上部鲜质量、地下部鲜质量、地下部干质量、株高和主茎高均较大,可作为首选指标,植株干质量、地上部干质量、主根长和出苗速率均较小,可作为辅助指标综合判断品种的耐盐能力.200个品种(系)在不同盐胁迫浓度下均可分成高度耐盐型、耐盐型、盐敏感型和高度盐敏感型4组.随盐胁迫强度加大,耐盐品种数量下降,而盐敏感品种数量上升.部分品种在低、中、高盐胁迫强度下表现出统一性(均耐盐或均敏感);部分品种存在差异性,即低胁迫强度下表现耐盐性而在高胁迫强度下表现盐敏感性.     

Comparisons of cadmium tolerance and accumulation at seedling stage in wheat varieties grown in different decades in China

Cadmium (Cd) tolerance and accumulation in wheat varieties were investigated at seedling stage under a controlled environmental condition. The 46 leading wheat varieties cultivated from the 1950s to 2000s in China were treated at the three-leaf stage with a 50 μM CdCl₂ solution for 24 days. Growth and photosynthesis parameters were measured and the Cd-tolerance index (ratio of a given parameter under Cd treatment to that of the control) was determined. Cd accumulation in shoots and roots and Cd translocation were also determined. It was found that Cd tolerance and accumulation of these wheat varieties varied over the different decades. Principal components analysis (PCA) showed that wheat varieties in 1950s and 1980s were tolerant while varieties from the last decade were sensitive to Cd stress. Wheat varieties in 1960s and 1970s were particularly tolerant to Cd stress for the parameters of shoot height, secondary root numbers, net photosynthesis and transpiration rate while the varieties in the 1990s were sensitive to Cd stress for shoot dry weight and root dry weight. Comparing each decade to the average Cd translocation ratio from the roots to the shoots for the whole period, the varieties from the 1950s and 1960s had a higher translocation ratio, while varieties in the 1970s were below that average. Varieties from the 1980s to 2000s showed an average translocation ratio. Using cluster analysis (CA), Shannongfu 63, Yangmai 1 and Yangmai 158 were the most Cd-tolerant varieties in which Cd translocation ratio were low, and Yumai 18 and Huaimai 20 were the most Cd-sensitive varieties in which Cd translocation ratio were high. The results indicating that wheat varieties from different decades were different in Cd tolerance and accumulation, and could be useful for breeding wheat for Cd stress tolerance.     

Constitutive and aluminium-induced patterns of phenolic compounds in two maize varieties differing in aluminium tolerance

Aluminium tolerance in maize is mainly due to more efficient Al exclusion. Nonetheless, even in tolerant varieties Al can gain access into the cells. Detoxification by binding to strong organic ligands should therefore play a role also in plants with high Al exclusion capacity. To test this hypothesis in this study the concentrations of soluble, free and bound, phenolics were analyzed in roots of two maize varieties differing in Al tolerance. Exposure for 24 h to 50 μM Al in nutrient solution strongly inhibited root elongation in the sensitive variety 16 × 36, but not in the Al-tolerant variety Cateto. Cateto accumulated about half the concentration of Al in roots than 16 × 36 (analysis performed after root desorption with citrate). Roots of Al-tolerant Cateto contained higher concentrations of caffeic acid, catechol and catechin than roots of the sensitive variety. Exposure to Al induced the accumulation of taxifolin in roots of both varieties. However, Al-tolerant Cateto accumulated about twice the concentration than Al-sensitive 16 × 36 of this pentahydroxyfavonol. The molar ratio for phenolics with catecholate groups to Al was about unity in roots of Cateto, while in those of 16 × 36 the ratio was ten times lower. Both the fact that these phenolics are strong ligands for Al and their high antioxidant and antiradical activity suggest that these compounds may provide protection against the Al fraction that is able to surpass the exclusion mechanisms operating in the tolerant maize variety.     

Distribution patterns of available P and K in rape rhizosphere in relation to genotypic difference

Plant genotypic differences in available K and P distribution at root-soil interface were studied with five rape varieties, which were 890206 (Brassica. campestris), Indian Mustard (Brassica juncea), Luzhoujinhuang (Brassica juncea), K-100 (Brassica juncea) and Xinongchangjiao (Brassica napus). The significant depletion of P and K was observed in the rhizosphere of all five rape varieties. The depletion rate of available P was in the range of 28–44% and the order followed: 890206 > Indian Mustard > K-100 > Xinongchangjiao > Luzhoujinhuang. The depletion area of available P occurred within 3–5 mm from root surface; and a wider depletion range was observed in the rhizosphere of the variety tolerant to low nutrient supply as compared to that of sensitive varieties. The depletion range of available K within the 10 mm from the root surface with the rate ranging from 31% to 48% and was wider than that of available P. The variety 890206 showed the highest absorption capability. Xinongchangjiao and Luzhoujinhuang showed higher depletion rate than that of K-100 and Indian Mustard, indicating that high-yield varieties of 890206 and Xinongchangjiao had higher absorption capacity of K than that of low-yield variety K-100. The results indicated that K depletion rate in the rhizosphere of rape was higher than that of cotton and was similar to that of two K enrichment plant–tobacco and alligator weed (Alternanthern Philoxeroides Mart. Griseb).     

花生苗期耐盐性评价及耐盐指标筛选

评价鉴选耐盐品种对于盐碱地花生生产具有重要意义。采用盆栽试验,设置不同盐胁迫浓度,对200个花生品种(系)萌发至幼苗期通过出苗速度、植株形态和生物量等指标进行耐盐性系统评价。结果表明,随盐胁迫浓度的增加,出苗时间延长,植株形态建成抑制加重,物质积累减少。鉴选花生品种耐盐性强弱的适宜盐胁迫浓度为0.30%—0.45%,超过此浓度不能出苗。地上部形态和生物量可作为耐盐评价的首选指标,主根长和出苗速度可作为辅助指标用以判断花生品种的综合耐盐能力。200个品种(系)在不同盐胁迫浓度下均可分成高度耐盐型、耐盐型、盐敏感型和高度盐敏感型4组。耐盐品种数量随盐胁迫强度加大而下降,盐敏感品种数量则上升。0.15%浓度下200个品种全部出苗,4个类型品种数分别占供试材料的29.0%、39.0%、27.5%和4.5%;0.30%浓度下185个品种出苗,4个类型品种数分别占供试材料的5.5%、34.5%、23.5%和29.0%;0.45%浓度下107个品种出苗,4个类型品种数分别占供试材料的5.5%、5.5%、20.0%和22.5%。14个品种在各盐浓度胁迫下均表现耐性,10个品种在各盐浓度胁迫下均表现敏感,为花生耐盐机理研究及生产应用提供了不同类型材料。     

耐盐水稻种质资源的筛选

用0.8% NaCl溶液和国际水稻所水稻耐盐性9级评价方法对前人选留的38份水稻耐盐种质资源(用0.5%NaCl盐土筛选)进行了重复筛选,以期获得极端耐盐水稻种质资源.同时研究了不同盐浓度对水稻发芽率的影响.结果表明:高浓度盐溶液显著抑制水稻种子的发芽率,但品种间有较大差异;筛选出苗期极端耐盐品种6份,其中1级1份,2级4份,3级1份;极端耐盐水稻品种苗高的增长受高浓度盐溶液抑制,但品种间差异不显著;而盐溶液对极端耐盐水稻品种出叶速率的影响因品种而异.     

Physiological responses of Tunisian grapevine varieties to bicarbonate-induced iron deficiency

Plants are frequently submitted to iron deficiency when growing on calcareous soils, which contain high concentrations of bicarbonate. The purpose of this study was to investigate the variability of physiological responses of Tunisian grapevine varieties to bicarbonate-induced iron chlorosis. Vine woodcuttings of seven autochthonous Tunisian varieties (Khamri, Mahdaoui, Blan3, Saouadi, Arich Dressé, Beldi and Balta4), two rootstocks (140Ru and S.O.4), and an introduced table variety (Cardinal) were cultivated on inert sand for 2 months using a complete nutrient solution (20 microM Fe) that was either well supplied or not supplied with 10 mM HCO3-. Young leaves of plants cultivated on bicarbonate-enriched medium showed characteristic symptoms of iron chlorosis, although the intensity of the symptoms depended on the variety and the rootstock. Chlorosis score confirmed these observations since the most sensitive varieties showed the highest values. This variability in tolerance to iron deficiency was also displayed when analysing the physiological parameters (shoot length, plant dry weight, and chlorophyll concentration) and the iron contents in the 4th leaf. Analysis of morphological and physiological parameters showed three behaviour groups. The first one corresponded to tolerant varieties (Khamri, Mahdaoui, and the root-stock: 140Ru), the second included moderately tolerant vines (Saouadi, Arich Dressé, Blanc3, and the rootstock: S.O.4) and the third represented the sensitive ones (Balta4, Beldi, and Cardinal).     

Physiological response of wheat varieties to elevated atmospheric CO2 and low water supply levels

In the phytotron experiment, the effect of elevated atmospheric CO₂ (EC, 750 μmol mol^?1) on the drought tolerance was studied in two winter varieties (Mv Mambo, tolerant; Mv Regiment, moderately tolerant) and in one spring variety of wheat (Lona, sensitive to drought). Changes in net photosynthetic rate (P _N), stomatal conductance, transpiration, wateruse efficiency, effective quantum yield of photosystem II, and activities of glutathione reductase (GR), glutathione-Stransferase (GST), guaiacol peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX) were monitored during water withdrawal. Drought caused a faster decline of P _N at EC, leading to the lower assimilation rates under severe drought compared with ambient CO₂ (NC). In the sensitive variety, P _N remained high for a longer period at EC. The growth at EC resulted in a more relaxed activation level of the antioxidant enzyme system in all three varieties, with very low activities of GR, GST, APX, and POD. The similar, low values were due to decreases in the varieties which had higher ambient values. A parallel increase of CAT was, however, recorded in two varieties. As the decline in P _N was faster at EC under drought but there was no change in the rate of electron transport compared to NC values, a higher level of oxidative stress was induced. This triggered a more pronounced, general response in the antioxidant enzyme system at EC, leading to very high activities of APX, CAT, and GST in all three varieties. The results indicated that EC had generally favourable effects on the development and stress tolerance of plants, although bigger foliage made the plants more prone to the water loss. The relaxation of the defence mechanisms increased potentially the risk of damage due to the higher level of oxidative stress at EC under severe drought compared with NC.     

Physiological performance of two contrasting rice varieties under water stress

Two rice varieties PR-115 and Super-7 were imposed to water stress and different physiological traits were monitored to evaluate the performance of these varieties under drought. Under water stress condition although the relative water content, osmotic potential, chlorophyll content, photosynthesis rate, carbon discrimination and biomass decreased in both the varieties however, the reduction was more pronounced in Super-7 variety. Oryzanol a trans-ester of ferulic acid functions as antioxidant and it increased along with total phenolic and anthocyanin content in both the varieties under drought stress. However, gallic acid, 4 hydroxy benzoic acid, syringic acid and chlorogenic acid showed differential pattern in both of the varieties under water limiting conditions. Under drought, grain yield was penalized by 17 and 54% in PR-115 and Super-7 varieties, respectively in comparison to watered plants. Super-7 variety showed pronounced electrolyte leakage and MDA enhancement under water stress condition. High non photochemical quenching and reduction in Y(NO) and Y(II) indicated balanced energy management in tolerant PR-115 variety. The studies showed that PR-115 is a drought tolerant variety while Super-7 is drought sensitive in nature.     

P32 distribution in phosphorus fractions of phosphorus-sensitive and -tolerant soybeans

Summary P³² incorporation into various phosphorylated compounds of roots, stems, and leaves was studied with P-sensitive (Clark) and P-tolerant (L9) varieties of soybeans. With increasing P concentration in the pretreatment solution, more P³² was incorporated into the inorganic phosphate fraction, mainly in which form P seemed to be transported from roots to leaves. P³² percentage of inorganic phosphate also showed a continuous decrease with absorption time. The rate of phosphorylation, as manifested by the ratio of P³² percentage or specific activities of the organic acid-soluble P to that of inorganic-P, was higher with the P-tolerant variety than with the P-sensitive one. The rate of RNA synthesis also seemed to be higher with the tolerant variety. Paper No. 7082, Scientific Journal Series, Minnesota Agricultural Experiment Station.     

22Na influx is significantly lower in salt tolerant groundnut (Arachis hypogaea) varieties

Distinct varieties differing in salt tolerance were initially identified from two separate green house experiments using two systems; solution as well as soil culture. The first screening involved a diverse group of 27 cultivars. Several physiological traits; Chlorophyll Stability Index (CSI), Salt Tolerance Index (STI) and ion content were determined to screen the cultivars for differences in salt tolerance using solution culture in the first experiment. A set of six varieties (three tolerant and three susceptible) were selected from this experiment and then subjected again to salt stress adopting a natural soil system in the second experiment which involved a screening approach essentially similar to that of the first experiment. In the third experiment using two distinct cultivars differing in salt tolerance selected from experiment II, ²²Na influx rate was determined in the root and shoot at the end of a 24 h salt imposition in Hoagland’s nutrient system containing 180 KBq of ²²Na. The results suggested that there were distinct differences in ²²Na influx rate into root and concurrently in the shoot. The salt tolerant Spanish improved and one of the moderately tolerant Trombay variety TAG 24, showed good regulation of ²²Na influx resulting in low ²²Na concentration. The salt susceptible variety JSP39 had nearly 7–8 fold higher root ²²Na content as compared to the tolerant and moderately tolerant cultivars. The results have highlighted the importance of Na exclusion as an important determinant of salt tolerance in groundnut.     

陆地棉萌发至三叶期不同生育阶段耐盐特性

摘要：土壤盐渍化是一个世界性的资源问题和生态问题,盐分胁迫几乎会影响棉花所有重要的生命活动, 造成棉花的减产或其他不利影响。本试验通过对14个品棉花种（系）的萌发期耐盐性、芽期耐盐性以及两个耐盐性不同的品种（系）三叶期的耐盐性,对棉花不同时期的耐盐特性进行了研究,结果表明：棉花在不同生育阶段耐盐能力是不同的,棉花盐敏感时间是在萌发出苗期。随着生育期的延长,棉花的耐盐性是逐步提高的,进行耐盐性鉴定的最佳时期在是萌发出苗期。在三叶期,随着盐分浓度的提高,棉花苗受伤害程度逐渐增加,盐敏感性品种受伤害大于耐盐品种,在同样盐浓度胁迫条件下,耐盐性不同的品种各部分受到的伤害不同,耐盐品种（系）中棉所35受伤害程度由高到低依次为：真叶>上胚轴>子叶;盐敏感品种（系）Tamcot CAB-CS受伤害程度由高到低依次为：真叶> 子叶>上胚轴。     

外源水杨酸预处理对高温胁迫下白菜耐热性和光合特性的影响

采用叶片喷施水杨酸（SA）的方法,研究高温胁迫下外源SA预处理对‘夏帝’和‘苏州青’2个白菜品种耐热性和光合特性的影响。结果表明：SA预处理降低了高温胁迫下白菜叶片的电解质渗透率、MDA和脯氨酸含量以及净光合速率（R）,从而缓解高温对质膜的过氧化伤害,并且通过提高可溶性糖和可溶性蛋白含量、SOD和POD活性来适应高温环境;然而SA预处理对气孔导度（GS）和蒸腾速率（功的影响在2个品种中变化相反,而对胞间CO2浓度（G）的影响差异不大。与耐热性较强的白菜品种‘夏帝’相比,耐热性较弱的品种‘苏州青’经SA预处理对缓解高温胁迫的影响效果更为明显。     

不同花生品种(系)萌发期抗旱性鉴定评价

以发芽势、发芽率、发芽指数、胚根长、胚轴长、幼芽长、幼苗鲜/干重的相对值为指标,通过PEG6000模拟干旱处理,对15份花生品种(系)进行萌发期抗旱性鉴定。结果显示,胁迫处理初期,低浓度处理对种子萌发有促进作用,随着胁迫处理时间的延长及胁迫处理浓度的升高,种子萌发受抑制程度增强。除相对幼芽长以外,其他相对指标都和品种综合抗旱能力呈极显著正相关。通过隶属函数法结合抗旱分级标准筛选出泰花4号、徐花13号、泰0125、泰0005等4份抗旱品种,泰花5号、濮花28、中花16等3份中抗材料,表明该方法可以作为一种快速、简便的鉴定花生萌发期抗旱性的方法。     

Adaptability to drought in sugar beet cultivars

The effects of NaCl and polyethylene glycol (PEG) on superoxide dismutase (SOD) and peroxidase (P) activities, lipid peroxidation (LP) and proline content in seeds and leaves of drought tolerant (FC-506 and MS-100) and drought sensitive (MS-612 and MS-13) sugar beet cultivars were examined. After PEG and NaCl treatment in tolerant cultivars both in seeds and leaves SOD activity mainly increased, though P activity increased only in leaves of tolerant cultivars. In drought sensitive cultivars the decrease of SOD and P activity was mostly observed. LP increased in seeds and leaves of all examined cultivars. The proline content increased in the leaves of examined cultivars and was significantly higher in drought tolerant plants. On the other hand, in the seeds only slight increase in proline content was found. The results obtained indicated that drought tolerance could be correlated with high proline content and enzymatic defense against lipid peroxidation.