安吉毛竹林生长季CO_2浓度的时空特征

试验采用7层CO2廓线系统对安吉毛竹林生长季(5—10月)CO2浓度进行连续监测,同时结合同步CO2通量和气象资料,分析安吉毛竹林CO2浓度的时空动态和CO2储存通量的变化特征。结果表明:安吉毛竹林生长季CO2浓度表现出明显的日变化、季节变化和垂直梯度。日尺度上,CO2浓度在日出前后达到最大,随后出现一段快速释放过程,午后达到全天最低值;整个生长季尺度上,7月份林间CO2浓度最低(368.1μmol·mol-1),10月份最高(402.6μmol·mol-1);垂直梯度方面,林冠上层CO2垂直梯度明显小于林冠层和林冠下层;林冠层和林冠上层垂直梯度变化趋势较为一致,白天均为正值,呈现为碳汇,夜间为负值,呈现为碳源,林冠下层CO2梯度全天均为负值。利用廓线系统计算毛竹林CO2储存通量,半小时和日尺度上,CO2储存通量分别占净生态系统CO2交换量(NEE)的25.5%和8.9%,而在月尺度上仅占NEE的0.22%。因此,在半小时及日尺度上计算NEE时不可忽略CO2储存通量,而在计算月尺度及更长时间尺度NEE时可以忽略CO2储存通量。     

黄河口潮间盐沼湿地生长季净生态系统CO2交换特征及其影响因素

潮间盐沼湿地生物地球化学过程独特,生态系统CO2交换存在着极大的复杂性和不确定性。利用2012年黄河口潮间盐沼湿地生态系统生长季(4—10月)连续的涡度相关观测数据,分析了潮间盐沼湿地的净生态系统CO2交换(NEE)、总初级生产力(GPP)和生态系统呼吸(Reco)的变化特征及其影响因素。结果表明:生长季,生态系统NEE具有明显的日变化和季节变化。日尺度上,表现为白天CO2净吸收,夜间CO2净释放,NEE日平均值为-0.38 g CO2m-2d-1;月尺度上,平均气温最高的7月生态系统释放CO2最多(15.16 g C/m2),6月生态系统吸收CO2最多(25.07 g C/m2)。潮间盐沼湿地生态系统的CO2交换受到光合有效辐射(PAR)、土壤温度(Ts)、土壤含水量(SWC)和潮汐淹水的共同影响。白天NEE主要受控于PAR,且生态系统表观初始光能利用率(α)和最大光合速率(NEEsat)分别在6月和5月达到最大值,分别为(0.0086±0.0019)μmol CO2μmol-1光子和(4.79±1.52)μmol CO2m-2s-1。夜间NEE随Ts呈指数增加趋势,生态系统呼吸的温度敏感性(Q10)为1.33,且SWC越高,Q10值越大。研究典型晴天(6月19日—6月25日)表明,潮汐淹水增强了生态系统白天对CO2的吸收,同时也增强了夜间CO2释放,研究时段内,潮汐淹水使生态系统净CO2吸收增加了0.76 g CO2m-2d-1。整个生长季,黄河口潮间盐沼湿地生态系统表现为CO2的汇,NEE为-22.28 g C/m2(其中,吸收118.34 g C/m2,释放96.28 g C/m2)。研究结果利于对潮间盐沼湿地源汇功能和影响机制的进一步认识与研究。     

发菜(Nostoc flagelliforme)培养条件的研究

研究了光照强度、日供水次数、CO2浓度和培养基成分对发菜生长的影响。结果显示,中度光强(114μmol.m-2.s-1)下发菜生长最快;发菜的生长基本同供水次数成正相关系;CO2浓度的升高并没有显著促进发菜生长,低光条件下(57μmol.m-2.s-1),高浓度的CO2(2800μL/L)抑制了发菜的生长;用BG11培养的发菜生物量的增长显著高于用BG110培养的;BG11培养基中K+和CO32-的缺失并没有显著影响发菜的生长。     

二氧化碳储存通量对森林生态系统碳收支的影响

涡度相关系统观测高度以下的CO₂储存通量对准确评价森林生态系统与大气间净CO₂交换量（NEE）有着重要的影响.本研究以长白山阔叶红松林为研究对象,利用2003年的涡度相关观测数据以及CO₂浓度廓线数据,分析了CO₂储存通量的变化规律及其对碳收支过程的影响.结果表明：涡度相关观测高度以下的CO₂储存通量具有典型的日变化特征,其最大变化量出现在大气稳定与不稳定层结转换期.利用涡度相关系统观测的单点CO₂浓度变化方法与利用CO₂浓度廓线方法计算的CO₂储存通量差异不显著.忽略CO₂储存通量,在半小时尺度上会造成对夜间和白天的NEE分别低估25%和19%,在日和年尺度上,会对NEE低估10%和25%;忽略CO₂储存通量,会低估Michaelis-Menten光响应方程及Lloyd-Taylor呼吸方程的参数,并且对表观初始量子效率α和参考呼吸R_ref的低估最大;忽略CO₂储存通量,在半小时、日及年尺度上,均会对总光合作用（GPP）和生态系统呼吸（R_e）低估约20%.     

黄河小浪底人工混交林生长季能量平衡特征

利用涡度相关系统和小气候梯度观测系统,对黄河小浪底人工混交林2012年生长季(5—9月)各能量通量进行了连续观测,分析了该生态系统能量平衡各项的变化特征,讨论了能量闭合状况。结果表明:潜热通量、感热通量和土壤热通量均与净辐射有类似的日变化特征。各项的绝对值大小表现为净辐射潜热通量感热通量土壤热通量。受日照时数的影响,5—9月能量平衡各项正值的日持续时间逐渐缩短。生长季,净辐射、感热通量和土壤热通量在6月份最大,最大值分别为418.5、231.4和12.5 MJ m-2month-1);潜热通量在7月份达到最大,最大值为320.8 MJ m-2month-1)。潜热通量、感热通量和土壤热通量占净辐射的比例分别在0.48—0.62、0.15—0.55、0.02—0.05之间。人工混交林生长季的能量分配主要以潜热通量和感热通量为主,且潜热通量为感热通量的2倍。波文比呈单峰曲线:6月最大,8月最小。黄河小浪底人工混交林生长季全天能量闭合度为79%。其中,白天闭合程度较高(81%),夜晚较低(41%)。本研究站点存在21%的能量不闭合。其原因可能与通量源区面积不匹配、忽略冠层热储存、湍流能的相位差等有关。     

温带针阔混交林土壤碳氮气体通量的主控因子与耦合关系

中高纬度森林地区由于气候条件变化剧烈,土壤温室气体排放量的估算存在很大的不确定性,并且不同碳氮气体通量的主控因子与耦合关系尚不明确。以长白山温带针阔混交林为研究对象,采用静态箱-气相色谱法连续4a(2005—2009年)测定土壤二氧化碳(CO2)、甲烷(CH4)和氧化亚氮(N2O)净交换通量以及温度、水分等相关环境因子。研究结果表明:温带针阔混交林土壤整体上表现为CO2和N2O的排放源和CH4的吸收汇。土壤CH4、CO2和N2O通量的年均值分别为-1.3 kg CH4hm-2a-1、15102.2 kg CO2hm-2a-1和6.13 kg N2O hm-2a-1。土壤CO2通量呈现明显的季节性规律,主要受土壤温度的影响,水分次之;土壤CH4通量的季节变化不明显,与土壤水分显著正相关;土壤N2O通量季节变化与土壤CO2通量相似,与土壤水分、温度显著正相关。土壤CO2通量和CH4通量不存在任何类型的耦合关系,与N2O通量也不存在耦合关系;土壤CH4和N2O通量之间表现为消长型耦合关系。这项研究显示温带针阔混交林土壤碳氮气体通量主要受环境因子驱动,不同气体通量产生与消耗之间存在复杂的耦合关系,下一步研究需要深入探讨环境变化对其耦合关系的影响以及内在的生物驱动机制。     

黄河三角洲芦苇湿地生态系统碳、水热通量特征

利用涡度相关法对黄河三角洲芦苇湿地生态系统进行了连续两年的通量观测,对2009—2010年生长季芦苇湿地的净生态系统碳交换量(NEE),感热通量(Hs)和潜热通量(LE)数据进行了分析。结果表明,在日尺度上,芦苇湿地NEE日变化特征表现为两个CO2吸收高峰,分别出现在11:00和16:00左右,其特点是在午间出现了碳交换通量的降低。CO2吸收的日最大值在两个生长季出现的时间有所不同,分别出现在2009年7月(-0.30 mg CO2m-2s-1)和2010年6月(-0.37 mg CO2m-2s-1)。CO2排放的日最大值两个生长季均出现在9月,分别为0.19和0.25 mg CO2m-2s-1。Hs和LE的日动态均为单峰型,极值都出现在中午前后,生长季生态系统的能量消耗主要以潜热为主,且在日尺度上,热通量和NEE有显著的负相关关系。在季节尺度上,芦苇湿地生长季具有明显的碳汇功能,2009年生长季生态系统白天固定354.63 g CO2/m2,同时期夜间释放159.24 g CO2/m2,净CO2吸收量为-195.39 g CO2/m2。2009年整个生长季生态系统总初级生产力(GPP)为-651.13 g CO2/m2,生态系统呼吸(Re)为455.74 g CO2/m2,系统表现为碳汇。路径分析表明:光合有效辐射(PAR)显著影响NEE的日动态(R2=0.46—0.84),而NEE的季节动态主要受土壤温度的影响,降水和PAR的影响次之。     

植被光合呼吸模型在长白山温带阔叶红松林的优化及验证

植被光合呼吸模型(VPRM)关键参数的确定和优化是准确计算生态系统净CO_2交换(NEE)的基础。利用中国通量观测研究联盟(China FLUX)长白山站温带阔叶红松林2005年的通量观测资料,对VPRM的4个参数(最大光能利用率ε_0、光照为半饱和条件下光合有效辐射值PAR0和呼吸参数(α、β))进行优化,并使用2006年的观测资料对参数优化前后的模拟结果进行评估。结果表明:参数优化后,VPRM能够较好地模拟长白山地区2006年植物生长季NEE的变化。对30min NEE模拟的平均误差为-1.81μmol m~(-2)s~(-1),相关系数为0.72,模拟NEE平均日变化的峰值约为观测值的91%,相关系数为0.97。但在植物非生长季模型对森林NEE的模拟效果较差。模型模拟30min NEE的平均误差为0.39μmol m~(-2)s~(-1),相关系数仅为0.10,并且模拟低估NEE平均日变化白天吸收峰值约82%,日变化模拟值与观测值的相关系数为0.50。通过分析不同天气个例,发现模型可以较好地模拟晴天条件下NEE的变化,而对阴雨天NEE的模拟误差较大。该研究有利于提高VPRM模型对温带落叶阔叶林NEE的模拟能力,对进一步改进区域陆地NEE的模拟具有重要意义。     

川西贡嘎山峨眉冷杉成熟林生态系统CO2通量特征

成熟森林的碳收支对陆地生态系统碳循环研究具有重要意义。目前,我国关于西南亚高山暗针叶林成熟林碳通量的研究还相对较少,尚不明确对碳循环的作用。以涡度相关技术为基础,对川西贡嘎山东坡峨眉冷杉成熟林生态系统尺度的CO_2通量进行长期定位观测。利用2015年6月至2016年5月观测数据,分析了峨眉冷杉成熟林净生态系统CO_2交换量(NEE)、生态系统呼吸(Re)和总生态系统生产力(GPP)的季节变异特征及其源汇状况,并结合环境因子,分析CO_2通量的主要控制因子。结果表明:(1)峨眉冷杉成熟林NEE具有明显的日变化特征,呈现"U"形变化,白天为负值,夜间为正值,中午前后CO_2通量达到最大;各月间日平均NEE变化差异显著,NEE峰值最大出现在2015年6月(-0.64 mg CO_2m~(-2)s~(-1)),峰值最小出现在2016年1月(-0.08 mg CO_2m~(-2)s~(-1));日平均NEE由正值变为负值的时间夏季最早,冬季最晚,NEE由负值变为正值的时间冬季最早,夏季最晚。(2)峨眉冷杉成熟林NEE、Re和GPP具有明显的月变化。2015年6月和12月NEE分别达到最大值(-46.02 g C m~(-2)月~(-1))和最小值(-1.42 g C m~(-2)月~(-1));Re呈现单峰变化,最大和最小值分别出现在2015年6月(84.78 g C m~(-2)月~(-1))和2016年1月(12.82 g C m~(-2)月~(-1));GPP最大值和最小值分别出现在2015年6月(130.81 g C m~(-2)月~(-1))与2016年1月(16.15 g C m~(-2)月~(-1))。(3)空气温度(T_a)、5 cm土壤温度(T_(s5))和光合有效辐射(PAR)是影响峨眉冷杉成熟林CO_2通量的主要环境因子。T_a与CO_2通量呈指数相关(R~2=0.5283,P0.01);白天CO_2通量与PAR显著相关(R~2=0.4373,P0.01);夜晚CO_2通量与T_(s5)显著相关(R~2=0.4717,P0.01)。(4)全年NEE、Re和GPP分别为-241.87、564.81 g C m~(-2)和806.68 g C m~(-2),表明川西贡嘎山峨眉冷杉成熟林具有较强的碳汇功能。     

鼎湖山针阔叶混交林冠层下方CO2通量及其环境响应

精确估算典型森林生态系统冠层下方CO₂通量（Fcb）对验证陆地生态系统碳平衡模型具有重要意义。采用开路涡度相关法对鼎湖山针阔叶混交林Fcb进行定位测定,根据1周年数据分析Fcb及其对环境要素的响应特征,结果表明：(1) 白天Fcb呈下降趋势表明地表植被全年具有光合能力,但总体上地表植被和土壤表现为CO2排放源;(2) Van’t Hoff方程、Arrhenius方程和Lloyd-Taylor方程均可以较好反映土壤温度(Ts)与Fcb的关系,其中仅Lloyd-Talor方程能够反映温度因子敏感性指标Q10随温度的变异性特征;(3) Lloyd-Talor方程模拟的Fcb完全由Ts控制,而连乘模型由Ts和土壤水分(Ms)控制,可以反映水热条件的综合影响,对Fcb具有更强的拟合能力;(4)在Ms较大时连乘模型对Fcb的估算高于Lloyd-Talor方程,反之在干旱时段连乘模型模拟结果低于Lloyd-Talor方程,表明当存在水分胁迫时,Ms可以成为影响Fcb的主导因子;(5) 2003年鼎湖山针阔叶混交林Fcb总量((787.4±296.8)gCm^-2a^-1)比静态箱-气相色谱法测得的土壤呼吸偏低17%。与箱式法相比,涡度相关法通量测定结果普遍存在偏低估算现象。     

A comparison of field methods for measuring soil carbon dioxide evolution: Experiments and simulation

Three widely used methods for measuring total soil CO₂ evolution are evaluated, including the dynamic CO₂ absorption method, the static CO₂ absorption method and the closed chamber method. The study covers laboratory experiments. numerical experiments with a simulation model and field measurements. The results are used to perform an error analysis. The aim of this error analysis is to indicate the impact of each method on the CO₂ dynamics during the measurement, and to select the most suitable method for frequent field usage.Laboratory experiments and simulation results show that the dynamic CO₂ absorption method has the potential to absorb all CO₂ evolving at the soil surface. The results also prove that the method has only a minor impact on the CO₂ concentration-depth gradient and the CO₂ efflux. The static CO₂ absorption method underestimates the soil CO₂ evolution, because the absorption velocity is too low, due to slow diffusion processes. Measurements with the closed-chamber method are based on an increasing concentration with time under a closed cover. However, the accumulation of gas alters the concentration gradient in the soil profile and thus causes a rapidly decreasing efflux during the measurement. A commonly used mathematical procedure, which corrects for the altered concentration gradient, does not yield the exact surface efflux, because the effect of increasing storage in the soil profile is not incorporated. Field measurements of CO₂ evolution, using the closed-chamber method and the dynamic CO₂ absorption method confirm the trends that have been predicted by the simulation model. The results of this study indicate that the dynamic CO₂ absorption method is accurate. As it is cheap and simple, it is suitable for the study of temporal and spatial dynamics of CO₂ evolution from the soil.     

Examination of the method for measuring soil respiration in cultivated land: Effect of carbon dioxide concentration on soil respiration

An acceleration of soil respiration with decreasing CO₂ concentration was suggested in the field measurements. The result supporrs that obtained in laboratory experiments in our previous study. The CO₂ concentrations in a chamber of the alkali absorption method (the AA-method) were about 150–250 parts/10⁶ lower than that in the atmosphere (about 350 parts/10⁶), while those observed in the open-flow IRGA method (the OF-method) were nearly equal to the soil surface CO₂ levels. The AA-method at such low CO₂ levels in the chamber appears to overestimate the soil respiration. Our results showed that the rates obtained by the AA-method were about twice as large as those by the OF-method in field and laboratory measurements. This finding has important consequences with respect to the validity of the existing data obtained by the AA-method and the estimation of changes in the terrestrial carbon flow with elevated CO₂     

用于超声造影的微囊藻气囊提取新方法

气囊是一种由蛋白质外壳包裹气体形成的纳米级细胞结构,常见于蓝藻和嗜盐古菌中.气囊中包含的气体在超声波作用下,能够散射声波并产生谐波信号而增强信噪比,具备成为新型超声造影剂的潜质.但目前提取气囊的方法主要是传统的高渗裂解法,该操作过程烦琐、得率低,不适用于气囊的大规模提取.针对这些技术瓶颈,文中建立了一种快速、高效的微囊...     

A comparison of stigmatellin conformations, free and bound to the photosynthetic reaction center and the cytochrome bc1 complex

We describe in detail the conformations of the inhibitor stigmatellin in its free form and bound to the ubiquinone-reducing (Q(B)) site of the reaction center and to the ubiquinol-oxidizing (Q(o)) site of the cytochrome bc(1) complex. We present here the first structures of a stereochemically correct stigmatellin in complexes with a bacterial reaction center and the yeast cytochrome bc1 complex. The conformations of the inhibitor bound to the two enzymes are not the same. We focus on the orientations of the stigmatellin side-chain relative to the chromone head group, and on the interaction of the stigmatellin side-chain with these membrane protein complexes. The different conformations of stigmatellin found illustrate the structural variability of the Q sites, which are affected by the same inhibitor. The free rotation about the chi1 dihedral angle is an essential factor for allowing stigmatellin to bind in both the reaction center and the cytochrome bc1 pocket.     

不同取样方式在稻田节肢动物采集中的效率评估

水稻是我国主要粮食作物,每年都会因虫害造成大量的经济损失,为了挽回害虫造成的损失,必须对害虫进行防治。田间节肢动物群落调查是评价害虫防治效果的重要依据,取样方式对节肢动物群落调查的准确性具有重要的影响。另外,对转基因作物对稻田生物多样性安全性进行评价时,取样方式对多样性评价的准确性也具有重要的影响。本文采用吸虫器法、盆拍法和马氏网诱集法3种取样方式进行稻田节肢动物调查,并评估不同取样方式的采集效率。得到的结果有:1.采集到的节肢动物物种数:马氏网诱集法吸虫器法盆拍法;2.采集的节肢动物数量:盆拍法吸虫器法马氏网诱集法;3.吸虫器取样法在调查叶蝉科、秆蝇科、茧蜂科、姬蜂科、金小蜂科、缘腹细蜂科、蕈蚋科时,取样效率较高;4.盆拍取样法在调查叶蝉科、瘿蚊科、微蛛亚科、跳蛛科、狼蛛科、猫蛛科、弹尾虫目、飞虱科时,取样效率较高;5.马氏网诱集法在调查缟蝇科和毛蠓科时取样效率较高。马氏网诱集法善于采集具有飞行能力的节肢动物;吸虫器法对不同习性的节肢动物采集效果均较高;盆拍法适合采集活动于水稻基部的节肢动物。     

A comparison and optimization of methods and factors affecting the transformation of Escherichia coli

DNA manipulation routinely requires competent bacteria that can be made using one of numerous methods. To determine the best methods, we compared four commonly used chemical methods (DMSO, MgCl₂–CaCl₂, CaCl₂ and Hanahan''s methods) on frequently used Escherichia coli (E. coli) strains: DH5α, XL-1 Blue, SCS110, JM109, TOP10 and BL21-(DE3)-PLysS. Hanahan''s method was found to be most effective for DH5α, XL-1 Blue and JM109 strains (P<0.05), whilst the CaCl₂ method was best for SCS110, TOP10 and BL21 strains (P<0.05). The use of SOB (super optimal broth) over LB [Luria–Bertani (broth)] growth media was found to enhance the competency of XL-1 Blue (P<0.05), dampened JM109′s competency (P<0.05), and had no effect on the other strains (P>0.05). We found no significant differences between using 45 or 90 s heat shock across all the six strains (P>0.05). Through further optimization by means of concentrating the aliquots, we were able to get further increases in transformation efficiencies. Based on the optimized parameters and methods, these common laboratory E. coli strains attained high levels of TrE (transformation efficiency), thus facilitating the production of highly efficient and cost-effective competent bacteria.     

Amount of nitrogen fixed by forage,pasture and grain legumes in Cyprus,estimated by the A-value and a modified difference method

The increasing need for protein at low cost has created a need to evaluate the biological nitrogen fixing potential of legumes in Cyprus. In field studies which were conducted over the growing years of 1982–3 and 1983–4, legumes which are traditionally grown in the country were evaluated for dry matter and nitrogen yield and biological nitrogen fixation (BNF). The legumes studied were medic (Medicago truncatula Gearth), ochrus vetch (Lathyrus ochrus L.), bitter vetch (Vicia ervilia L.) and faba bean (Vicia faba L. var major) in the first year and in addition chickpea (Cicer arietinum L.), woollypod vetch (Vicia dasycarpa Ten.) and tickbean (Vicia faba L. var minor) in the second year. Using the A-value method with barley and oats as reference crops, nitrogen (N) fixed by the various legumes in the first year was 30–50% and from 55–67% of total N yield for the two reference crops, respectively. In the second year the estimates of N fixed ranged from 70 to 80% with similar results obtained for the two reference crops barley and ryegrass. However, in the second year chickpea, which had limited nodulation, fixed only 40% of its N yield. Estimates of nitrogen from the atmosphere (Ndfa) obtained by the difference method (DM) were 10 to 14% lower than those from the A-value method. These results were obtained after correcting for the amount of N derived from the applied fertilizer. The two methods were highly correlated (r=0.98) for estimates of amount of BNF. The rates of N₂ fixation of uninoculated legumes which are nodulated by the indigenous populations of Rhizobium in Cyprus are comparable to those of legumes inoculated with selected strains of Rhizobium in other countries. An exception was the amount of N fixed by chickpea. The appearance of the first nodules at late stages of growth may be the reason for the low BNF of this crop.     

Variable alteration of regional tissue oxygen pressure in rat hippocampus by acute swimming exercise

One of the events in the brain is an increasing cerebral blood flow during exercise. The tissue oxygen level may be increased because blood flow correlates with tissue oxygen level. However, it is little known whether the tissue oxygen pressure in hippocampal region (Hip-pO₂) will be affected by exercise.     

Dipalmitoylphosphatidylcholine membranes modified with carotenoid pigment lutein: Experiment versus Monte Carlo simulation study of the membrane organization

Organization of bilayer lipid membrane formed with dipalmitoylphosphatidylcholine and containing a xanthophyll pigment lutein was studied by both the Monte Carlo simulation and UV-Vis absorption spectroscopy. The simulations were based on ten-state Pink model. The proposed model consisted of two monolayers represented by a two-dimensional triangular lattice with vacancies. The orientation and aggregation state of lutein, obtained from the analysis of the spectroscopic measurements, were used to calibrate intermolecular interactions in the model. In accordance with the experimental data, the model allows two orientations of lutein molecules: one spanning the membrane and the other parallel to its plane. The influence of the intermolecular interactions on the main phase transition as well as on the aggregation of lutein molecules is discussed. The analysis of the model enables us to learn about molecular mechanisms that govern the effects of lutein on the membrane properties as well as the effects of the lipid matrix on lutein organization in the membrane. A concept is discussed according to which increasing domination of parallel lutein orientation, observed at high temperatures, can protect the membrane against penetration by water molecules and reactive oxygen species and against loss of the membrane compactness, especially in the regions of oxidized acyl chains.     

Solution structure of an atypical WW domain in a novel beta-clam-like dimeric form

The WW domain is known as one of the smallest protein modules with a triple-stranded beta-sheet fold. Here, we present the solution structure of the second WW domain from the mouse salvador homolog 1 protein. This WW domain forms a homodimer with a beta-clam-like motif, as evidenced by size exclusion chromatography, analytical ultracentrifugation and NMR spectroscopy. While typical WW domains are believed to function as monomeric modules that recognize proline-rich sequences, by using conserved aromatic and hydrophobic residues that are solvent-exposed on the surface of the beta-sheet, this WW domain buries these residues in the dimer interface.