烏飯樹(shù)和藍(lán)莓對(duì)不同土壤pH值的生理反應(yīng)

李晴晴，魯珊珊，張紅，楊艷，肖家欣

（安徽師范大學(xué)生命科學(xué)學(xué)院/安徽省重要生物資源保護(hù)與利用研究重點(diǎn)實(shí)驗(yàn)室，安徽蕪湖241000）

烏飯樹(shù)和藍(lán)莓對(duì)不同土壤pH值的生理反應(yīng)

李晴晴，魯珊珊，張紅，楊艷，肖家欣*1

（安徽師范大學(xué)生命科學(xué)學(xué)院/安徽省重要生物資源保護(hù)與利用研究重點(diǎn)實(shí)驗(yàn)室，安徽蕪湖241000）

采用盆栽砂培法，研究3種不同的土壤pH值（4.2、5.2和6.2）對(duì)烏飯樹(shù)（Vaccinium bracteatum Thunb.）和兔眼藍(lán)莓（Vaccinium ashei Reade）品種“燦爛”生長(zhǎng)及相關(guān)生理指標(biāo)的影響。結(jié)果表明：烏飯樹(shù)和藍(lán)莓的生物量、葉綠素含量及根系活力均在土壤pH 6.2時(shí)最低，而其生物量與根系活力分別在pH 5.2和pH 4.2時(shí)最高。烏飯樹(shù)和藍(lán)莓根和葉部的可溶性糖及可溶性蛋白質(zhì)含量均在pH 6.2時(shí)最高，分別在pH 5.2和pH 4.2時(shí)最低。在土壤pH 5.2時(shí)，烏飯樹(shù)的根、莖、葉中P、Fe和Zn含量較高；當(dāng)土壤pH 4.2時(shí)，藍(lán)莓根部P和Fe含量、莖部Fe含量及葉部Fe和Zn含量均相對(duì)較高。烏飯樹(shù)和藍(lán)莓根或葉的POD和CAT活性及MDA含量均在pH 6.2時(shí)最高，而其最低值分別出現(xiàn)在pH 5.2和pH 4.2時(shí)。綜上，烏飯樹(shù)和藍(lán)莓分別在土壤pH 5.2和pH 4.2時(shí)生長(zhǎng)最好，pH 6.2的土壤環(huán)境對(duì)二者生長(zhǎng)均有抑制作用，尤其以藍(lán)莓較為明顯。

土壤pH；烏飯樹(shù)；藍(lán)莓；生理反應(yīng)

SummaryBlueberry(Vaccinium spp.)is well known for its rich anthocyanins and other bioactive compounds,which contributes to prevent from cardiovascular disease and other chronic illnesses.Thereby,many countries including China have attached great importance to the economic value and development prospect of blueberry.However,the main root of blueberry tree is not obvious but exceedingly slender,intertwined and lack of root hair.Meanwhile,blueberry prefers acidic even strongly acidic soil,and its most suitable pH value of culture is from 3.8 to 5.0.As the soil pH value of cultivating blueberry is generally higher than 5.0,large amounts of sulfur powder are applied to reduce the soil pH value. Wufanshu(Vaccinium bracteatum Thunb.),belonging to the same family and genus with blueberry,is a stock with developed root system,strong barren tolerance and extensively ecological adaptability,which has been cultivated as a kind of economic or landscaping plant species.There are many examples of grafting and improving native blueberry with Wufanshu scion in folk.Grafting blueberry with Wufanshu is well worth popularizing in blueberry cultivation at least in the Yangtze River area of China.However,few studies have been reported on the effects of soil pH values on the growth of different Vaccinium plants.

To evaluate the effects of different soil pH values on the growth and relative physiological indices of Wufanshu(V. bracteatum)and rabbiteye blueberry(Vaccinium ashei Reade)cv.Brightwell plants,three different soil pH values(4.2,5.2 and 6.2)were applied to the 9-month-old plants of Wufanshu and rabbiteye blueberry under greenhouse conditions.Plants were grown in the medium of V(sand)∶V(perlite)=1∶1 and irrigated every 2 days with half-strength Hoagland’s No.2 nutrient solutions with different pH values(4.2,5.2,and 6.2).Leaves,stems and roots were separately sampled after 187 days to determine their relative physiological indices.

The results showed that the biomass,chlorophyll content and root activity of Wufanshu and blueberry were the lowest at pH 6.2 level,but the maxima of the biomass and root activity in Wufanshu and blueberry were found at pH 5.2 and 4.2 levels,respectively.The soluble sugar and protein contents of Wufanshu and blueberry were the highest at pH 6.2,and the lowest ones were detected at pH 5.2 and 4.2,respectively.The contents of phosphorus(P),ferrum(Fe)and zinc(Zn)in the roots,stems and leaves of Wufanshu were the highest at pH 5.2 level;however,the contents of P and Fe in the roots,Fe in the stems,and Fe and Zn in the leaves of blueberry were the highest at pH 4.2 level.The activities of peroxidase(POD)and catalase(CAT),and malondialdehyde(MDA)content of Wufanshu and blueberry were the highest at pH 6.2 level,while the minima of them were presented at pH 5.2 and 4.2,respectively.

These results indicate that pH 5.2 and pH 4.2 are the optimal soil pH values for the growth of Wufanshu and blueberry plants,respectively;however,soil pH 6.2 will inhibit their growth,especially the blueberry.

烏飯樹(shù)（Vaccinium bracteatum Thunb.）為杜鵑花科越橘屬灌木或小喬木，耐瘠薄，較耐寒，為淺根性樹(shù)種，喜酸性環(huán)境，在土壤pH 4.5～6.6的黃紅壤或紅壤上生長(zhǎng)良好，是南方酸性紅壤區(qū)一種良好的水土保持植物，也是酸性紅壤土的指示植物之一[1]。藍(lán)莓（Vaccinium spp.）亦為杜鵑花科越橘屬灌木，其果實(shí)富含花青苷，低糖，低脂肪，抗氧化能力強(qiáng)，具有增強(qiáng)免疫力、保護(hù)視力和心臟功能的獨(dú)特療效，營(yíng)養(yǎng)價(jià)值高，被國(guó)際糧農(nóng)組織列為人類5大健康食品之一[2]。藍(lán)莓屬淺根性樹(shù)種，無(wú)根毛，最適土壤pH值為3.8～5.0，然而天然土壤pH值往往高于5.0，因此許多地區(qū)采用施用硫磺粉的方法來(lái)降低土壤pH值以適應(yīng)其栽培需求，這不僅限制了藍(lán)莓的廣泛推廣，還增加了栽培成本[3-5]。近年來(lái)，我國(guó)藍(lán)莓產(chǎn)業(yè)發(fā)展勢(shì)頭強(qiáng)勁，由北向南已推廣到江南、華南等地。2015年全國(guó)超過(guò)20個(gè)省市區(qū)開(kāi)展了藍(lán)莓種植，總面積約3萬(wàn)hm2，產(chǎn)量約2.5萬(wàn)t[6]。藍(lán)莓通常分為3大類：高叢藍(lán)莓（北高叢、南高叢和半高叢藍(lán)莓）、兔眼藍(lán)莓及矮叢藍(lán)莓，其中在長(zhǎng)江流域廣泛引種栽培的大多是南高叢藍(lán)莓和兔眼藍(lán)莓[7]。土壤pH值過(guò)高容易引起藍(lán)莓黃化等異常癥狀，一般認(rèn)為當(dāng)土壤pH值高于5.2時(shí)，土壤中的自由態(tài)鐵會(huì)與有機(jī)物質(zhì)作用，生成不能被藍(lán)莓根系吸收的絡(luò)合物，從而引起藍(lán)莓植株缺鐵黃化等癥狀[8]。

生產(chǎn)上除了施用硫磺粉降低土壤pH值來(lái)適應(yīng)藍(lán)莓栽培的需求外，曾有研究者以烏飯樹(shù)為砧木嫁接藍(lán)莓并獲得成功[9-11]。烏飯樹(shù)和藍(lán)莓親緣關(guān)系較近，而且烏飯樹(shù)對(duì)土壤的適應(yīng)性相對(duì)較強(qiáng)，在我國(guó)南北各地均有分布[12]，因此，烏飯樹(shù)作為藍(lán)莓嫁接的砧木具有廣泛的應(yīng)用前景。烏飯樹(shù)和藍(lán)莓雖然都喜歡酸性土壤條件，但二者對(duì)土壤適應(yīng)性的生理差異尚不明確。因此，本研究以南方普遍栽培的烏飯樹(shù)和兔眼藍(lán)莓品種“燦爛”為試驗(yàn)材料，研究二者對(duì)不同土壤pH值的生理反應(yīng)差異，明確它們最適生長(zhǎng)的土壤pH值，為進(jìn)一步提高藍(lán)莓栽培的適應(yīng)性提供理論與實(shí)踐依據(jù)。

1 材料與方法

1.1 試驗(yàn)材料

試驗(yàn)基質(zhì)為V（石英砂）∶V（珍珠巖）=1∶1的混合物，清洗干凈后，取出自然晾干，裝入3 L不透光塑料盆中，每盆約3.0 kg。試驗(yàn)用的兔眼藍(lán)莓（Vaccinium ashei Reade）品種“燦爛”和烏飯樹(shù)（Vaccinium bracteatum Thunb.）苗分別購(gòu)自安徽徽王農(nóng)業(yè)有限公司和江蘇省宜興市南燭烏飯樹(shù)生態(tài)園。

1.2 試驗(yàn)設(shè)計(jì)

2015年4月10日，將大小相對(duì)一致的9個(gè)月齡的藍(lán)莓苗和烏飯樹(shù)苗移栽入裝有3.0 kg試驗(yàn)基質(zhì)的塑料盆（盆上口內(nèi)徑22 cm、盆底內(nèi)徑16 cm、盆高18 cm）中，然后放置于溫室大棚內(nèi)，每隔1 d澆1次水（200 mL/盆，pH 5.0）。待植株萌發(fā)出新葉后，開(kāi)始澆灌1/4濃度Hoagland營(yíng)養(yǎng)液（pH 5.0），每隔2 d澆灌1次，澆灌2次后洗鹽1次。待植物長(zhǎng)到7～8片新葉（約7周）時(shí)澆灌1/2濃度Hoagland營(yíng)養(yǎng)液，同時(shí)進(jìn)行不同pH處理。設(shè)置3個(gè)pH梯度：pH 4.2、pH 5.2和pH 6.2，分別用1 mol/L鹽酸和氫氧化鈉在pH檢測(cè)儀下調(diào)配。采用完全隨機(jī)分組，共6個(gè)處理，每個(gè)處理6盆，每盆定植1株，共36盆。2015年10月17日分別取葉片、根系及莖進(jìn)行相關(guān)生理指標(biāo)的測(cè)定。

1.3 分析方法

植株采收后將部分樣品置于105℃烘箱中殺青15 min，之后置于70℃條件下烘至恒量，用稱質(zhì)量法測(cè)定單株根和莖的干質(zhì)量。葉綠素含量采用乙醇丙酮浸提法測(cè)定，可溶性糖與可溶性蛋白質(zhì)含量分別采用蒽酮比色法和G-250比色法測(cè)定[10]。分別將藍(lán)莓和烏飯樹(shù)的根、莖、葉干樣用不銹鋼電動(dòng)粉碎機(jī)粉碎，然后采用硝酸-高氯酸（體積比為4∶1）消煮法消煮，用Optimal 2100 DV電感耦合等離子體發(fā)射光譜儀（Pekin-Elmer公司，美國(guó)）測(cè)定消化液中P、 Fe、Zn和Mg含量[11]。過(guò)氧化物酶（peroxidase,POD）活性采用愈創(chuàng)木酚法測(cè)定，過(guò)氧化氫酶（catalase, CAT）活性采用紫外吸收法測(cè)定，丙二醛（malondialdehyde,MDA）含量采用硫代巴比妥酸比色法測(cè)定[13]。

1.4 數(shù)據(jù)處理

所獲得的數(shù)據(jù)采用SPSS 20.0及Excel 2007進(jìn)行統(tǒng)計(jì)分析。其中，利用SPSS 20.0軟件的單向方差分析（one-way analysis of variance）進(jìn)行不同處理間的差異顯著性檢驗(yàn)，采用鄧肯法進(jìn)行多重比較。

2 結(jié)果與分析

2.1pH值對(duì)烏飯樹(shù)和藍(lán)莓生物量、葉綠素含量與根系活力的影響

當(dāng)營(yíng)養(yǎng)液pH值為4.2和5.2時(shí)，烏飯樹(shù)和藍(lán)莓葉片的葉綠素含量均顯著高于pH 6.2時(shí)的含量；烏飯樹(shù)根和莖的干質(zhì)量以及根系活力在營(yíng)養(yǎng)液pH 5.2時(shí)最高；而藍(lán)莓的莖干質(zhì)量及根系活力在pH 4.2時(shí)最高，且顯著高于pH 5.2和pH 6.2時(shí)的對(duì)應(yīng)值，其根干質(zhì)量在pH 4.2和pH 5.2時(shí)的差異無(wú)統(tǒng)計(jì)學(xué)意義，且均高于pH 6.2時(shí)的對(duì)應(yīng)值（表1）。

表1pH值對(duì)烏飯樹(shù)和藍(lán)莓生物量、葉綠素含量與根系活力的影響Table1Effect of pH values on biomass,chlorophyll content and root activity of Wufanshu and blueberry

2.2pH值對(duì)烏飯樹(shù)和藍(lán)莓可溶性糖與可溶性蛋白質(zhì)含量的影響

烏飯樹(shù)根和葉部的可溶性糖與可溶性蛋白質(zhì)含量均以pH 6.2時(shí)最高，pH 4.2次之，pH 5.2時(shí)最低；而藍(lán)莓根部可溶性糖與可溶性蛋白質(zhì)含量均以pH 4.2時(shí)最低，pH 5.2居中，pH 6.2時(shí)最高（圖1）。

圖1pH值對(duì)烏飯樹(shù)和藍(lán)莓可溶性糖與可溶性蛋白質(zhì)含量的影響Fig.1Effect of pH values on soluble sugar and soluble protein contents in Wufanshu and blueberry

2.3pH值對(duì)烏飯樹(shù)和藍(lán)莓礦質(zhì)元素含量的影響

從圖2可以看出：在3個(gè)不同pH值處理下，烏飯樹(shù)根、莖和葉部P、Fe和Zn含量均以pH 5.2時(shí)最高，且除pH 4.2與pH 5.2時(shí)葉部Zn含量在統(tǒng)計(jì)學(xué)上無(wú)顯著差異外，其他都呈現(xiàn)顯著性差異；而pH 5.2時(shí)烏飯樹(shù)根、莖和葉部Mg含量顯著低于pH 4.2和pH 6.2時(shí)的含量，且除根部Mg含量在pH 4.2與pH 5.2時(shí)無(wú)顯著差異外，其余都差異顯著。當(dāng)pH 4.2時(shí)，藍(lán)莓根部P和Fe、莖部Fe及葉部Fe和Zn含量均顯著高于pH 5.2和pH 6.2時(shí)的含量，除pH 4.2和pH 5.2時(shí)藍(lán)莓根部Zn含量無(wú)顯著差異外，其余均顯著高于pH 6.2時(shí)的含量；而pH 4.2時(shí)藍(lán)莓根、莖和葉部Mg含量均顯著低于pH 5.2和pH 6.2時(shí)的含量；此外，3個(gè)不同pH值之間的藍(lán)莓莖和葉部P含量，以及莖部Zn含量均無(wú)顯著差異。

圖2pH值對(duì)烏飯樹(shù)和藍(lán)莓根、莖及葉片中P、Mg、Fe、Zn元素含量的影響Fig.2Effect of pH values on P,Mg,Fe and Zn contents in the roots,stems and leaves of Wufanshu and blueberry

2.4pH值對(duì)烏飯樹(shù)和藍(lán)莓CAT和POD活性與MDA含量的影響

在pH 5.2時(shí)，烏飯樹(shù)根和葉的CAT和POD活性以及MDA含量均顯著低于pH 4.2與pH 6.2時(shí)的值，且葉片CAT活性在pH 4.2與pH 5.2時(shí)差異不顯著，其余均呈現(xiàn)顯著差異；在pH 4.2時(shí)，藍(lán)莓根部CAT活性及MDA含量均顯著低于pH 6.2和pH 5.2時(shí)的對(duì)應(yīng)值，而在pH 4.2和pH 5.2時(shí)藍(lán)莓根部POD活性均低于pH 6.2時(shí)的值；烏飯樹(shù)和藍(lán)莓根或葉的POD、CAT活性及MDA含量均以pH 6.2時(shí)最高（圖3）。

圖3pH值對(duì)烏飯樹(shù)和藍(lán)莓CAT和POD活性與MDA含量的影響Fig.3Effect of pH values on CAT and POD activities and MDA content in Wufanshu and blueberry

3 討論

土壤pH值，即土壤酸堿度是一個(gè)重要的土壤理化指標(biāo)，對(duì)植物生長(zhǎng)、土壤微生物活動(dòng)、養(yǎng)分轉(zhuǎn)化以及土壤肥力等均有明顯的影響[14]。適宜植物生長(zhǎng)的土壤pH值一般有一定的范圍，只有在合適的pH值范圍內(nèi)，植物才能充分吸收和利用養(yǎng)分，生長(zhǎng)良好。土壤pH值過(guò)高或過(guò)低對(duì)植物而言都是一種環(huán)境脅迫，會(huì)影響植物的生長(zhǎng)發(fā)育和生理代謝過(guò)程，如培養(yǎng)基pH過(guò)低或過(guò)高會(huì)抑制非生根藍(lán)莓組培苗的生長(zhǎng)[15]。當(dāng)然，植物對(duì)逆境的變化程度也具有一定的適應(yīng)能力[16]。

有研究認(rèn)為，藍(lán)莓正常生長(zhǎng)對(duì)土壤酸堿度的要求極為嚴(yán)格，其最適土壤pH值為4.0～4.8[17]。鄉(xiāng)土樹(shù)種烏飯樹(shù)與藍(lán)莓的親緣關(guān)系較近（為杜鵑花科越橘屬灌木或小喬木），而且該樹(shù)種對(duì)土壤的適應(yīng)性強(qiáng)，在中國(guó)南北各地均有分布[12]。本研究結(jié)果顯示，烏飯樹(shù)根系活力及根、莖干質(zhì)量均以pH 5.2時(shí)最高，而藍(lán)莓則以pH 4.2時(shí)最高，二者葉綠素含量均以pH 6.2時(shí)最低?？梢?jiàn)，烏飯樹(shù)與藍(lán)莓均適宜在酸性環(huán)境中生長(zhǎng)，pH 6.2的土壤環(huán)境對(duì)二者植株生長(zhǎng)均有一定的抑制作用。這不同于柑橘或其他樹(shù)種生長(zhǎng)的最適pH水平為6.0左右[18]。

可溶性糖和可溶性蛋白質(zhì)均為滲透調(diào)節(jié)物質(zhì)。在逆境條件下，植物體內(nèi)的滲透調(diào)節(jié)物質(zhì)往往會(huì)升高，從而增加細(xì)胞液濃度，降低水勢(shì)，促進(jìn)植物吸收水分，以抵抗不良環(huán)境對(duì)植物細(xì)胞的傷害。本研究表明，烏飯樹(shù)的根、葉和藍(lán)莓根部的可溶性糖與可溶性蛋白質(zhì)含量分別在pH 5.2和pH 4.2時(shí)最低，并均在pH 6.2時(shí)最高?？梢?jiàn)，pH 6.2時(shí)二者的滲透調(diào)節(jié)物質(zhì)積累明顯，說(shuō)明烏飯樹(shù)和藍(lán)莓植株已處于一種脅迫環(huán)境。這與前人在杜鵑[19]和大豆[20]上的研究結(jié)果類似，即當(dāng)土壤pH值處于脅迫水平時(shí)，植株體內(nèi)的滲透調(diào)節(jié)物質(zhì)含量明顯上升。另一方面，pH 5.2和pH 4.2分別是烏飯樹(shù)和藍(lán)莓較合適的土壤pH值，因而在合適的pH條件下，其滲透調(diào)節(jié)物質(zhì)積累較少，這與上述根系活力和生物量結(jié)果亦相吻合。

本研究結(jié)果還顯示，在不同pH值條件下，烏飯樹(shù)各部位P、Fe和Zn含量均在pH 5.2時(shí)最高，而藍(lán)莓則在pH 4.2時(shí)對(duì)P、Fe和Zn的吸收量最多。這表明pH 5.2和pH 4.2分別是烏飯樹(shù)和藍(lán)莓較合適的土壤pH條件，在此條件下最有利于植株對(duì)P、Fe和Zn等礦質(zhì)元素的吸收與轉(zhuǎn)運(yùn)。而Mg含量在烏飯樹(shù)和藍(lán)莓的最適pH條件下均為最低，這與李亞?wèn)|等[21]的研究結(jié)果類似，但與曹增強(qiáng)等[15]的研究結(jié)果略有不同。曹增強(qiáng)等[15]認(rèn)為，培養(yǎng)基pH過(guò)高（6.0、6.5和7.0）或過(guò)低（pH 4.5）均會(huì)抑制藍(lán)莓組培苗對(duì)P、K、Zn和Mg等元素的吸收。這可能與藍(lán)莓苗的培養(yǎng)條件（非生根組培苗培養(yǎng)基培養(yǎng)與砂基培養(yǎng)）差異有關(guān)。一般認(rèn)為，在酸性環(huán)境下Mg的溶解度增加，容易淋失，植株因來(lái)不及吸收而造成Mg元素的缺乏。另一方面，礦質(zhì)元素之間還存在相互拮抗的關(guān)系，這也可能使Mg元素的吸收轉(zhuǎn)運(yùn)與其他元素不同[22]。因而，在實(shí)際生產(chǎn)中，在酸性土壤上栽培藍(lán)莓和烏飯樹(shù)要注意及時(shí)補(bǔ)充Mg肥，以防止缺Mg現(xiàn)象的發(fā)生。

植物在水分脅迫和重金屬污染等逆境條件下,體內(nèi)活性氧含量往往會(huì)上升,為防止過(guò)量的活性氧對(duì)植物細(xì)胞膜的傷害,植株體內(nèi)抗氧化酶活性也會(huì)明顯提高，從而使活性氧含量維持在較低水平[23-25]。過(guò)氧化氫酶（CAT）和過(guò)氧化物酶（POD）可以催化H2O2轉(zhuǎn)化為H2O。丙二醛（MDA）是膜脂過(guò)氧化反應(yīng)的產(chǎn)物，其含量變化在一定程度上反映了細(xì)胞受傷害的程度。本研究結(jié)果顯示，無(wú)論是烏飯樹(shù)還是藍(lán)莓，其根或葉的POD和CAT活性及MDA含量均在pH 6.2時(shí)最高。這與王明元等[26]的研究結(jié)果一致，即隨著基質(zhì)pH值的升高，枳幼苗受到的脅迫明顯加重，植株體內(nèi)的POD、CAT活性均明顯提高。可見(jiàn)，pH 6.2的土壤環(huán)境對(duì)烏飯樹(shù)和藍(lán)莓都造成了一定的逆境傷害，為應(yīng)對(duì)該環(huán)境脅迫，植物體內(nèi)的POD和CAT活性升高，抗氧化防御能力增強(qiáng)，但其生長(zhǎng)仍受到明顯抑制。在pH 5.2時(shí)，烏飯樹(shù)根或葉的CAT、POD活性及MDA含量均達(dá)到最低，而藍(lán)莓則在pH 4.2時(shí)最低。這進(jìn)一步說(shuō)明土壤pH 5.2對(duì)烏飯樹(shù)生長(zhǎng)較為合適，而pH 4.2的土壤條件對(duì)藍(lán)莓生長(zhǎng)較為合適。

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Physiological response to different soil pH values between Vaccinium bracteatum and Vaccinium ashei.

Journal of Zhejiang University(Agric.&Life Sci),2017,43(4):469-475

LI Qingqing,LU Shanshan,ZHANG Hong,YANG Yan,XIAO Jiaxin*(Anhui Provincial Key Laboratory of the Concervation and Exploitation of Biological Resources/College of Life Sciences,Anhui Normal University,Wuhu 241000,Anhui,China)

soil pH;Vaccinium bracteatum;Vaccinium ashei;physiological response

S 663；Q 945

A

10.3785/j.issn.1008-9209.2016.08.281

國(guó)家自然科學(xué)基金（31372014）；安徽省高等學(xué)校自然科學(xué)研究項(xiàng)目（KJ2016SD24）。

肖家欣（http://orcid.org/0000-0002-8427-6551），E-mail:xjx0930@163.com

（First author）：李晴晴（http://orcid.org/0000-0002-9957-8018），E-mail:lqq9201@163.com

2016-08-28；接受日期(Accepted)：2016-10-22