摘要:【目的】對(duì)切花月季彎梗表型性狀進(jìn)行分析和篩選,并綜合評(píng)價(jià)不同品種的彎梗特性,為矯正不同類型月季彎梗及促進(jìn)切花月季優(yōu)質(zhì)高效生產(chǎn)提供參考依據(jù)?!痉椒ā恳詼厥以耘嗟?0個(gè)切花月季品種為研究對(duì)象,統(tǒng)計(jì)不同品種的彎梗率;測(cè)定彎梗植株的花朵寬度、花朵高度、萼片下落距離等12個(gè)表型性狀,統(tǒng)計(jì)各指標(biāo)的變異系數(shù)(CV);采用主成分分析(PCA)和聚類分析等方法綜合評(píng)價(jià)10個(gè)切花月季品種的彎梗特性?!窘Y(jié)果】切花月季品種高盛的彎梗率最高,達(dá)16.19%,白荔枝彎梗率最小,僅3.44%。氣場(chǎng)和蘇醒的彎梗萼片下落后變化類型分別為3和2種,其余品種為1種;品種間花朵鮮重等10個(gè)表型性狀變異系數(shù)大于10.00%,各品種彎梗方向無(wú)特定方位。已測(cè)的12個(gè)表型性狀中有5對(duì)指標(biāo)間呈極顯著相關(guān)(Plt;0.01),5對(duì)指標(biāo)間呈顯著相關(guān)(Plt;0.05)。PCA結(jié)果表明,莖粗、最大花瓣厚度、花朵鮮重、花瓣數(shù)、花朵寬、花盤到第1臺(tái)葉距離、下落萼片鮮重和花梗高等8個(gè)性狀載荷量較大,是彎梗特性評(píng)價(jià)的關(guān)鍵指標(biāo)。通過隸屬函數(shù)法對(duì)綜合評(píng)價(jià)值(D)排序?yàn)楦呤t;浪漫沙灘gt;艾莎gt;夏日戀情gt;洛神gt;粉荔枝gt;蘇醒gt;氣場(chǎng)gt;朱麗葉gt;白荔枝。聚類分析結(jié)果顯示供試切花月季品種可分為三大類群,高盛、浪漫沙灘和艾莎具有突出的彎梗特性,D均大于0.650?!窘Y(jié)論】10個(gè)切花月季品種彎梗植株表型性狀變異較大,氣場(chǎng)和蘇醒彎梗萼片表型變化較多;篩選出莖粗、最大花瓣厚度、花朵鮮重、花瓣數(shù)、花朵寬、花盤到第1臺(tái)葉距離、下落萼片鮮重和花梗高等8個(gè)關(guān)鍵指標(biāo)用于月季彎梗特性評(píng)價(jià);獲得高盛、浪漫沙灘和艾莎3個(gè)彎梗特性較強(qiáng)的切花月季品種,可作為研究切花月季彎梗特性的材料。
關(guān)鍵詞:切花月季;表型性狀;彎梗特性;主成分分析;聚類分析;綜合評(píng)價(jià)
中圖分類號(hào):S685.12文獻(xiàn)標(biāo)志碼:A文章編號(hào):2095-1191(2024)09-2679-10
Comprehensive evaluation of cut rose bent peduncle characteris?tics based on principal component analysis and cluster analysis
YANG Jiao1,LUO Zhong-yuan2,TAO Jin-ting1,MENG Jing1,ZHAO Yan1*
(1College of Landscape and Horticulture,Yunnan Agricultural University,Kunming,Yunnan 650201,China;
2Yao’an County Rural Industry Development Center,Chuxiong,Yunnan 675300,China)
Abstract:【Objective】The purpose of this study was to analyze and screen the bent peduncle phenotypic traits of cut rose,and comprehensively evaluate the bent peduncle characteristics of different varieties of cut rose,so as to provide re-ference for correcting different types of cut rose bent peduncle and promoting the high quality and efficient production of cut rose.【Method】Ten cut rose varieties cultivated in greenhouse were studied,and the bent peduncle rate of different va-rieties was calculated.The 12 phenotypic characters such as flower width,flower height and sepal fall distance were deter-mined in bent peduncle plants,and the coefficient of variation(CV)of each index was counted.Principal component analysis(PCA)and cluster analysis were used to evaluate the bent peduncle characteristics of 10 cut rose varieties.【Re-sult】The cut rose variety Gotcha had the highest bent peduncle rate(16.19%),and the White O’Hara had the lowest bent peduncle rate(3.44%).There were 3 and 2 change types of Glamorous and Awakening after bent peduncle sepals fell re-spectively,and 1 in other varieties.The CV of 10 phenotypic traits,such as fresh flower weight,was greater than 10.00%,and there was no specific orientation in the bent peduncle direction of each variety.Among the 12 phenotypic traits tested,5 pairs were extremely significantly correlated with each other(Plt;0.01)and 5 pairs were significantly correlated with each other(Plt;0.05).The results of PCA showed that stem diameter,maximum petal thickness,flower fresh weight,petal number,flower width,distance from disk to the first leaf,fresh weight of falling sepal and height of peduncle were the 8 key indicators for evaluating the characteristics of the bent peduncle since they had large load.According to the mem-bership function method,the comprehensive evaluation value(D)was ranked as Gotchagt;Romantic Beachgt;Jumiliagt;Summer Lovegt;Goddess of the Luo Rivergt;Pink O’Haragt;Awakeninggt;Glamorous+gt;Julietgt;White O’Hara.The clus-ter analysis results showed that the cut rose varieties in the test could be divided into 3 groups,Gotcha,Romantic Beach and Jumilia had outstanding bent peduncle characteristics with D greater than 0.650.【Conclusion】The variation of pheno-typic characters of 10 cut rose varieties with bent peduncle plant is great.The phenotypes of sepals in Glamorous+and Awakening bent peduncle have more changes.Eight key indexes,including stem diameter,maximum petal thickness,flower fresh weight,petal number,flower width,distance from disk to the first leaf,fresh weight of falling sepal and pe-duncle height,are selected to evaluate the characteristics of rose bent peduncle.Three cut rose varieties with strong bent peduncle characteristics,Gotcha,Romantic Beach and Jumilia,are obtained,which can be used as materials to study the bent peduncle characteristics of cut rose.
Key words:cut rose;phenotypic traits;bent peduncle characteristics;principal component analysis;cluster analy-sis;comprehensive evaluation
Foundation items:National Natural Science Foundation of China(31660683);General Project of Yunnan Agricul-ture Joint Project(2301BD070001-032);General Project of Yunnan Fundamental Research Project(202201AT070256)
0引言
【研究意義】切花月季(Rosa hybrida L.)在世界花卉產(chǎn)業(yè)和國(guó)際鮮切花貿(mào)易中占據(jù)重要地位,位列四大鮮切花之首(黃小艷等,2023)。云南省是我國(guó)切花月季的主要產(chǎn)地之一(張海兵等,2021),昆明花拍中心公布2023年切花月季的產(chǎn)量為11億枝,產(chǎn)值達(dá)13億元。然而切花月季在種植中普遍存在彎?,F(xiàn)象(Bent peduncle phenomenon,BPP),BPP并非采后因水分脅迫引起的彎頸現(xiàn)象,而是一種生理性畸形(Zaccai et al.,2009;石力勻等,2018),嚴(yán)重的會(huì)失去經(jīng)濟(jì)價(jià)值(石力勻等,2020)。BPP發(fā)生率因品種而異,如單頭品種Mercedes的BPP發(fā)生率為10%~30%(Zaccai et al.,2009),Beast為10%~20%(Seo and Kim,2013),卡羅拉約為20%(石力勻等,2020),蜜桃雪山為15%~25%(Jing et al.,2020)。因此,分析和篩選切花月季彎梗表型性狀,綜合評(píng)價(jià)切花月季的彎梗特性,對(duì)矯正不同類型月季彎梗及促進(jìn)切花月季優(yōu)質(zhì)高效生產(chǎn)具有重要意義?!厩叭搜芯窟M(jìn)展】切花月季發(fā)生BPP時(shí),子房下方花梗會(huì)朝1個(gè)方向彎曲(角度為5°~90°),有1個(gè)特定萼片下落并發(fā)展成葉狀結(jié)構(gòu),萼片下落方向與花梗彎曲方向一致,特定萼片原本位置則出現(xiàn)萼瓣化現(xiàn)象?;ü澢糠殖0橛斜馄交F(xiàn)象,影響花器官發(fā)育,出現(xiàn)花梗變短、花瓣數(shù)和花朵鮮重減少、花朵變小等表型變化(Seo and Kim,2013;石力勻等,2020;王晗,2023)。表型性狀直接反映植物的外部特征,是植物多樣性評(píng)價(jià)的主要參考依據(jù)(張斌斌等,2021)。植物表型研究的關(guān)鍵是獲取高質(zhì)量并可重復(fù)的性狀數(shù)據(jù),進(jìn)而量化分析基因型和環(huán)境互作效應(yīng)及其對(duì)產(chǎn)量、質(zhì)量、抗逆等主要性狀的影響(Ribaut etal.,2010;Tes-ter and Langridge,2010)。在分析種質(zhì)和植株表型性狀差異時(shí),由于性狀指標(biāo)數(shù)量多且各指標(biāo)間具有相關(guān)性,難以用普通統(tǒng)計(jì)學(xué)方法對(duì)材料和性狀進(jìn)行綜合評(píng)價(jià)(曹婷婷等,2024)。因此,利用主成分分析(PCA)法對(duì)性狀指標(biāo)降維,處理性狀指標(biāo)間的重疊信息,提取少量獨(dú)立的綜合指標(biāo)可表達(dá)大部分信息(許小宛等,2019)。吳超等(2018)用相關(guān)性分析和PCA研究微型盆栽月季,篩選出花型、葉型、花色等5個(gè)重要性狀;過聰?shù)龋?019)對(duì)60個(gè)現(xiàn)代月季品種表型性狀進(jìn)行研究,通過PCA篩選出葉型和花型2個(gè)評(píng)價(jià)月季性狀的重要因子;Khaleghi和Khadivi(2020)利用PCA對(duì)大馬士革玫瑰的36個(gè)形態(tài)性狀進(jìn)行評(píng)價(jià),篩選重要的性狀,從而獲得優(yōu)質(zhì)材料;尹世華等(2021)對(duì)47個(gè)月季品種進(jìn)行表型多樣性分析及綜合評(píng)價(jià),篩選獲得綜合評(píng)分最高的月季品種甜蜜漂流;孫佩等(2022)通過聚類分析對(duì)119份月季品質(zhì)表型性狀進(jìn)行綜合評(píng)價(jià),獲得一些表型性狀好且環(huán)境適應(yīng)性強(qiáng)的月季品種;王莉飛等(2022)結(jié)合PCA和聚類分析對(duì)57份現(xiàn)代月季種質(zhì)資源的13個(gè)表型性狀進(jìn)行綜合評(píng)價(jià),篩選出皮刺、花頭數(shù)等5個(gè)性狀,可作為月季種質(zhì)評(píng)價(jià)的重要指標(biāo);吉乃喆等(2023)通過研究65個(gè)藤本月季品種的表型,利用PCA和聚類分析將藤本月季分為4組。目前,國(guó)內(nèi)外學(xué)者對(duì)切花月季彎梗的表型性狀及彎梗特性研究較少,大部分研究集中在彎梗植株激素等方面。Zaccai等(2009)通過研究Mercedes彎梗植株的萼片和花莖的內(nèi)源激素,結(jié)果發(fā)現(xiàn)彎梗與生長(zhǎng)素相關(guān),外源激素萘乙酸(NAA)和吲哚乙酸(IAA)能導(dǎo)致正常植株短暫彎梗,生長(zhǎng)素運(yùn)輸抑制劑(NPA)則能矯正彎梗植株。Seo和Kim(2013)通過研究切花月季Beast的花部形態(tài)、生長(zhǎng)和內(nèi)源激素水平,結(jié)果發(fā)現(xiàn)彎梗由花的同化產(chǎn)物不均衡分配和高IAA含量引起。Jing等(2020)通過分析月季品種蜜桃雪山花梗的轉(zhuǎn)錄譜和細(xì)胞結(jié)構(gòu),結(jié)果發(fā)現(xiàn)彎梗與生長(zhǎng)素、赤霉素和細(xì)胞分裂素有關(guān)?!颈狙芯壳腥朦c(diǎn)】我國(guó)的月季栽培歷史悠久,月季育種始于我國(guó),但目前我國(guó)月季育種存在新品種數(shù)量較少、品質(zhì)較差等問題(劉鑫穎等,2022)。遺傳多樣性是培育新品種的重要基礎(chǔ),對(duì)植物進(jìn)行表型性狀遺傳多樣性分析可為品種選育提供參考(王業(yè)舉等,2023,喬雨軒等,2024)。目前有關(guān)切花月季彎梗的研究存在測(cè)試表型性狀少及對(duì)彎梗特性的評(píng)價(jià)缺乏系統(tǒng)性等問題,而基于PCA和聚類分析的切花月季彎梗特性綜合評(píng)價(jià)尚未見研究報(bào)道?!緮M解決的關(guān)鍵問題】以10個(gè)切花月季品種彎梗植株為試驗(yàn)材料,測(cè)定植株花梗高、莖粗、彎梗植株的萼片下落距離等12個(gè)與表型相關(guān)的性狀指標(biāo),分析不同品種彎梗植株的差異性,通過表型性狀多樣性分析、PCA、聚類分析和隸屬函數(shù)法對(duì)其彎梗特性進(jìn)行綜合評(píng)價(jià),為矯正不同類型月季彎梗及促進(jìn)切花月季優(yōu)質(zhì)高效生產(chǎn)提供參考依據(jù)。
1材料與方法
1.1試驗(yàn)材料
試驗(yàn)地位于云南省楚雄州姚安縣的切花月季花卉大棚,栽培基質(zhì)體積比為珍珠巖∶椰糠∶蛭石=1∶8∶1,生長(zhǎng)溫度為15~30℃/10~15℃(晝/夜),相對(duì)濕度為75%~80%,光照時(shí)間為10~14 h,光照強(qiáng)度為35000~50000 lx。2023年2月種植扦插苗,之后選取長(zhǎng)勢(shì)和開放度相似的第4茬植株為試驗(yàn)材料,品種信息見表1。
1.2試驗(yàn)方法
于2023年9月中旬上午8:00—12:00測(cè)量表型性狀。參考李玲玲(2020)的五點(diǎn)采樣法,統(tǒng)計(jì)每個(gè)樣點(diǎn)的彎梗率。參考石力勻等(2020)的方法和LYT 1868—2010《植物新品種特異性、一致性、穩(wěn)定性測(cè)試指南—薔薇屬》,對(duì)正常植株和彎梗植株的形態(tài)進(jìn)行拍照;使用游標(biāo)卡尺測(cè)量彎梗植株的萼片下落距離、花盤到第1臺(tái)葉的距離、花朵寬度、花朵高度和莖粗;使用卷尺測(cè)量花梗高度;使用植物葉片厚度計(jì)測(cè)量最大花瓣厚度和下落萼片厚度;使用指南針和量角器分別測(cè)量彎梗方位和彎梗角度;使用精密電子天平稱量下落萼片鮮重和花朵鮮重,統(tǒng)計(jì)樣株的花瓣數(shù)量,每個(gè)品種重復(fù)10次。計(jì)算方差及變異系數(shù)(CV)。
CV(%)=S/ˉ(x)×100
式中,S表示標(biāo)準(zhǔn)差,ˉ(x)表示性狀平均值。1.3統(tǒng)計(jì)分析
使用PowerPoint 2019繪制圖片,采用Excel 2019、SPSS 26.0和Origin 2021進(jìn)行數(shù)據(jù)整理、統(tǒng)計(jì)分析和作圖。參考于碩等(2023)的方法繪制數(shù)據(jù)量化后的彎梗方位雷達(dá)圖。參考藺豆豆等(2021)、周玉杰等(2023)的評(píng)價(jià)及計(jì)算方法,對(duì)性狀指標(biāo)進(jìn)行相關(guān)分析、PCA和聚類分析。采用隸屬函數(shù)法綜合評(píng)價(jià)彎梗特性,根據(jù)綜合評(píng)價(jià)值(D)進(jìn)行聚類分析,度量標(biāo)準(zhǔn)采用平方歐式距離,并制出分類結(jié)果譜系圖。各品種綜合指標(biāo)隸屬函數(shù)值、權(quán)重和綜合評(píng)價(jià)計(jì)算方法如下:
式中,Ui表示i品種的第i指標(biāo)的隸屬函數(shù)值,Xi表示第i個(gè)綜合指標(biāo);Xmin表示第i個(gè)綜合指標(biāo)最小值;Xmax表示第i個(gè)綜合指標(biāo)最大值。Wi表示第i個(gè)綜合指標(biāo)在所有綜合指標(biāo)中重要程度即權(quán)重;Pi表示各基因型第i個(gè)綜合指標(biāo)貢獻(xiàn)率;D為各基因型由綜合評(píng)價(jià)所得的彎梗特性綜合評(píng)價(jià)值。
2結(jié)果與分析
2.1 10個(gè)切花月季品種的彎梗率和彎梗方位分析結(jié)果
由圖1-A可知,在10個(gè)切花月季品種中,高盛的彎梗率最高,達(dá)16.19%;夏日戀情次之,與高盛無(wú)顯著差異(Pgt;0.05,下同);白荔枝的彎梗率最小,僅為3.44%,白荔枝、粉荔枝、浪漫沙灘、蘇醒和朱麗葉5個(gè)品種間彎梗率無(wú)顯著差異,這5個(gè)品種的彎梗率顯著小于另外5個(gè)品種(Plt;0.05,下同)。由圖1-B可知,高盛、洛神和艾莎3個(gè)品種在西南方位的彎梗居多,分別為22.86%、20.83%和20.00%;蘇醒在東北和西北方位彎梗均為37.29%;白荔枝和粉荔枝東方位彎梗較多,分別為33.33%和27.27%;浪漫沙灘在東南方位彎梗占比為50.00%;朱麗葉在西北方位彎梗占比38.46%;夏日戀情和氣場(chǎng)彎梗向南方位居多,分別為25.71%和21.05%,說(shuō)明切花月季彎梗無(wú)特定方位,且與太陽(yáng)的方向無(wú)關(guān)。
2.2 10個(gè)切花月季品種正?;ㄅc彎梗花的表型分析結(jié)果
由圖2-A可知,與正常植株相比,彎梗植株的花梗由圓柱狀變?yōu)楸馄?,彎曲部位直徑變大,花頭變小。花萼5個(gè),有1個(gè)萼片下落,下落到花梗的部位發(fā)生彎曲,彎曲方向與下移萼片方向一致。大部分彎梗植株有1個(gè)變異萼片的位置被半瓣半萼或萼瓣?duì)畹钠鞴俅妫ò甑捷嗥霓D(zhuǎn)換均是發(fā)生在擴(kuò)大的葉狀結(jié)構(gòu)下方。由圖2-B可知,變異萼片下落后出現(xiàn)4種表型:無(wú)萼瓣?duì)钶嗥˙S)、下落后變異萼片原本著生的位置由1個(gè)萼瓣?duì)钊〈˙S1)、由2個(gè)萼瓣?duì)钊〈˙S2)、由3個(gè)萼瓣?duì)钇鞴偃〈˙S3)。其中氣場(chǎng)月季品種的萼片下落后有BS1、BS2和BS3等3種表型,蘇醒有BS和BS1等2種表型,其余8個(gè)品種僅出現(xiàn)1種表型(BS或BS1)。
進(jìn)一步對(duì)彎梗植株的12個(gè)性狀進(jìn)行統(tǒng)計(jì)分析,結(jié)果如表2所示。除花朵寬和下落萼片厚度2個(gè)性狀外,其余10個(gè)性狀CV均大于10.00%。品種間表型性狀變異性最大的是花朵鮮重,CV為89.42%,極差為16.73,平均值為3.78 g。萼片下落距離變異性次之,CV為79.33%,平均值為30.53 mm;萼片下落距離最大值為120.40 mm,最小值為1.50 mm,二者差異明顯。變異性最小的性狀為下落萼片厚度,CV為9.09%,平均值為0.33 mm。
2.3 10個(gè)切花月季品種彎梗性狀指標(biāo)間相關(guān)分析結(jié)果
對(duì)上述12個(gè)表型性狀指標(biāo)進(jìn)行相關(guān)性分析,結(jié)果(圖3)可知,有5對(duì)指標(biāo)間呈極顯著相關(guān)(rlt;0.01,下同),5對(duì)指標(biāo)間呈顯著相關(guān)(rlt;0.05,下同)。莖粗與下落萼片厚度、花朵高與花朵鮮重、下落萼片厚度與彎梗角度呈顯著正相關(guān);花朵鮮重與花瓣數(shù)、下落萼片厚度與花瓣數(shù)呈顯著負(fù)相關(guān)。莖粗與最大花瓣厚度和花朵鮮重、最大花瓣厚度與下落萼片厚度呈極顯著正相關(guān);花瓣數(shù)與莖粗和最大花瓣厚度均呈極顯著負(fù)相關(guān)。其中莖粗與花朵鮮重的相關(guān)系數(shù)最大,為0.89。
2.4 10個(gè)切花月季品種彎梗性狀指標(biāo)的PCA結(jié)果
進(jìn)一步用主成分降維法對(duì)12個(gè)指標(biāo)進(jìn)行PCA得到因子荷載矩陣,結(jié)果如表3所示。將累計(jì)方差貢獻(xiàn)率大于80%且特征值≥1作為判別條件,可提取4項(xiàng)主成分因子,即第一主成分(PC1)~第四主成分(PC4),累計(jì)貢獻(xiàn)率達(dá)90.645%,說(shuō)明4個(gè)主成分可以反映12個(gè)性狀90.645%的形狀特征。PC1的特征值為4.910,貢獻(xiàn)率為40.918%,主要反映月季彎梗的花朵和花莖的形態(tài)特征,其中莖粗、最大花瓣厚度、花朵鮮重和花瓣數(shù)的載荷量絕對(duì)值均超過0.800;PC2的特征值為2.859,貢獻(xiàn)率為23.822%,其載荷量絕對(duì)值最大的是花朵寬,為0.838;PC3的特征值為2.018,貢獻(xiàn)率為16.818%,載荷量絕對(duì)值高于0.800的是花盤到第1臺(tái)葉的距離;PC4的特征值為1.090,貢獻(xiàn)率為9.087%,載荷量絕對(duì)值高于0.500的性狀是下落萼片鮮重和花梗高,主要反映彎梗月季的萼片和花梗的特征情況。因此,莖粗、最大花瓣厚度、花朵鮮重、花朵寬、花瓣數(shù)、花盤到第1臺(tái)葉的距離、下落萼片鮮重和花梗高這8個(gè)表型性狀,能反映月季品種彎梗植株的花、莖和變形萼片部位的形態(tài)性狀基本特征,且其貢獻(xiàn)率與實(shí)際鑒別分類形態(tài)依據(jù)基本一致。因此,前4個(gè)主成分中載荷量絕對(duì)值最大的8個(gè)性狀指標(biāo)可作為研究切花月季彎梗特性的關(guān)鍵性指標(biāo)。
2.5 10個(gè)切花月季品種的彎梗特性隸屬函數(shù)綜合評(píng)價(jià)結(jié)果
綜合評(píng)價(jià)10個(gè)切花月季品種的彎梗特性,結(jié)果如表4所示,對(duì)于綜合指標(biāo)X1而言,浪漫沙灘的隸屬函數(shù)值最大,U1為1.000,而白荔枝的隸屬函數(shù)值最小,U1為0.000,說(shuō)明在X1下,浪漫沙灘表現(xiàn)出最強(qiáng)的彎梗特性,而白荔枝的彎梗特性最弱。計(jì)算出U1、U2、U3、U4的權(quán)重分別為0.451、0.263、0.186、0.100。按10個(gè)切花月季品種彎梗特性的D排序?yàn)楦呤t;浪漫沙灘gt;艾莎gt;夏日戀情gt;洛神gt;粉荔枝gt;蘇醒gt;氣場(chǎng)gt;朱麗葉gt;白荔枝。
2.6 10個(gè)切花月季品種彎梗植株聚類分析結(jié)果
根據(jù)D采用平均距離聯(lián)接法進(jìn)行聚類分析。結(jié)合表4和圖4可知,以平均歐式距離10為界限,供試切花月季品種可分為三大類群,第Ⅰ類群包括高盛、浪漫沙灘和艾莎,這3個(gè)品種D均大于0.650,具有較強(qiáng)的彎梗特性;第Ⅱ類群包括洛神、夏日戀情、朱麗葉、氣場(chǎng)、蘇醒和粉荔枝,這6個(gè)品種D為0.340~0.551,該類群切花月季品種的彎梗特性相對(duì)較弱;第Ⅲ類群為白荔枝,其D為0.193,該類群彎梗特性最弱,最不易發(fā)生彎梗。聚類分析結(jié)果與D排序基本一致。
3討論
莖稈質(zhì)地強(qiáng)健、挺直、無(wú)彎頸是決定鮮切花品質(zhì)的關(guān)鍵指標(biāo)(黃麗娟,2020),莖稈發(fā)生彎曲和傾斜會(huì)降低切花的觀賞價(jià)值和商品價(jià)值(石文波,2021)。植物花莖或花梗彎曲現(xiàn)象較為常見,如非洲菊的花莖在瓶插期會(huì)因花頭的重力(Ferrante etal.,2007)、脫落酸的調(diào)節(jié)(Ge etal.,2019)、莖中果膠和半纖維素的解離(Cheng et al.,2020)等原因出現(xiàn)彎曲。擬南芥在花蕾形成時(shí),淀粉體位移區(qū)域受重力影響,會(huì)導(dǎo)致整個(gè)花序軸彎曲(Weiseetal.,2000;Powers and Strader,2020)。切花月季花梗彎曲呈鉤狀會(huì)導(dǎo)致花不能正常開放,研究表明彎梗發(fā)生率與品種有關(guān)(Zaccai et al.,2009;石力勻等,2020)。本研究中10個(gè)切花月季品種的彎梗率為3.44%~16.19%,說(shuō)明不同品種間差異較明顯,與前人研究結(jié)果一致,且無(wú)固定彎梗方位。切花月季花器官和花梗發(fā)生改變是彎梗的典型特征,彎梗中萼片擴(kuò)大和變形呈現(xiàn)出葉狀結(jié)構(gòu),萼片不正常發(fā)育導(dǎo)致月季花朵畸形(Zaccai etal.,2009;Wang et al.,2019;Jinget al.,2020),降低切花月季商品價(jià)值。在月季中葉狀結(jié)構(gòu)取代花瓣、萼片、雄蕊、雌蕊等花器官的現(xiàn)象較為常見,如綠萼和Motrea的花器官轉(zhuǎn)化成畸形的葉狀結(jié)構(gòu)(Mor and Zieslin,1992;Chmelnitsky et al.,2003;Sim et al.,2004;Yan et al.,2016)。本研究發(fā)現(xiàn),供試品種彎梗植株均出現(xiàn)葉變和萼瓣化現(xiàn)象,有1個(gè)變異萼片類似葉子,發(fā)生高度不等的下落現(xiàn)象,在花蕾下方的花梗上發(fā)育,變異萼片所在部位變得扁平,所在方向與花梗彎曲方向一致。變異萼片本應(yīng)著生的位置沒有萼瓣化器官或被1、2、3個(gè)萼瓣化器官替代,氣場(chǎng)和蘇醒的彎梗萼片變化類型較多,可用作后期研究切花月季彎梗的重要材料。
表型性狀具有直觀、易識(shí)別的特點(diǎn),通過認(rèn)真細(xì)致地觀察可區(qū)分出不同的表型個(gè)體(肖滿秋等,2023)。表型性狀CV的大小充分反映材料的變異程度,CV大于10%時(shí),說(shuō)明材料間差異較大(Liu et al.,2004)。本研究中除花朵寬和下落萼片厚度性狀外,有10個(gè)性狀CV均大于10.00%,說(shuō)明本研究篩選的10個(gè)切花月季品種的形態(tài)多樣性較高,可作為后期研究切花月季彎梗植株的品種。性狀指標(biāo)相關(guān)分析是對(duì)2個(gè)或2個(gè)以上有相關(guān)性的變量進(jìn)行分析,從而確定性狀指標(biāo)之間的相關(guān)程度(尹世華等,2021)。本研究中12個(gè)表型性狀之間的相關(guān)性聯(lián)系較多,呈極顯著和顯著關(guān)系的性狀各有5對(duì),其中相關(guān)性聯(lián)系最大的是莖粗與花朵鮮重,說(shuō)明評(píng)價(jià)切花月季彎梗特性與篩選指標(biāo)的過程中,可著重考慮莖粗對(duì)花朵鮮重的影響。利用PCA對(duì)種質(zhì)表型性狀評(píng)價(jià)進(jìn)行探討,而PCA篩選出的性狀指標(biāo)可通過種質(zhì)的聚類分析分組情況進(jìn)行驗(yàn)證(翟藝蘭等,2023)。本研究對(duì)12個(gè)表型性狀進(jìn)行PCA,4個(gè)主成分累計(jì)貢獻(xiàn)率達(dá)到90.645%,選取載荷量絕對(duì)值最大的8個(gè)指標(biāo)用于評(píng)價(jià)月季彎梗特性,較石力勻等(2020)的研究性狀指標(biāo)個(gè)數(shù)較多。隸屬函數(shù)法能夠在多指標(biāo)測(cè)定的基礎(chǔ)上,通過計(jì)算供試指標(biāo)的D及隸屬函數(shù)值,根據(jù)D對(duì)供試材料進(jìn)行綜合評(píng)價(jià),避免各指標(biāo)因貢獻(xiàn)率不同產(chǎn)生的誤差(藺豆豆等,2021)。本研究對(duì)供試材料所測(cè)性狀指標(biāo)進(jìn)行綜合評(píng)價(jià),結(jié)果發(fā)現(xiàn)高盛彎梗特性最強(qiáng),白荔枝最不易發(fā)生BPP。根據(jù)D聚類分析將供試材料分為三大類群,其中高盛、浪漫沙灘和艾莎表型相似且彎梗特性較強(qiáng),與表型性狀調(diào)查結(jié)果基本一致。
此外,研究表明月季彎梗受基因和生長(zhǎng)素等內(nèi)部條件的共同調(diào)控,月季的A、B、C、D 4組基因分別控制萼片和花瓣、萼片和雄蕊、雄蕊和心皮、胚珠和心皮的發(fā)育,E組基因則參與了全部5輪花器官的發(fā)育,如果基因發(fā)生突變或異常表達(dá),花器官的形成將出現(xiàn)異常(Ditta et al.,2004;Litt and Kramer,2010;Wellmeretal.,2014;石力勻等,2018)。月季植株從營(yíng)養(yǎng)生長(zhǎng)到生殖生長(zhǎng)過程中激素不均衡分布,花器官發(fā)育的A、B類功能基因異常表達(dá)容易造成萼片葉變現(xiàn)象,引發(fā)花瓣萼瓣化(Yan et al.,2016;石力勻,2020;Jinget al.,2020)。月季的生長(zhǎng)素和赤霉素主要來(lái)源于花器官(Faragher etal.,1987),Zaccai等(2009)研究了單個(gè)月季品種彎梗植株的萼片變化和內(nèi)源激素,發(fā)現(xiàn)彎梗出現(xiàn)與生長(zhǎng)素有關(guān),在月季正常植株的花梗部注射生長(zhǎng)素,會(huì)出現(xiàn)跟畸形花類似的彎折現(xiàn)象。Jing等(2020)研究了月季花梗轉(zhuǎn)錄譜和細(xì)胞結(jié)構(gòu),認(rèn)為彎梗處調(diào)控生長(zhǎng)素的RhTMK1和RhIAA32基因表達(dá)水平較低,與細(xì)胞分裂素的調(diào)控基因RhARR3和RhARR8表達(dá)水平存在差異,同時(shí)赤霉素調(diào)控基因RhGA20-0x和RhGA2-ox的表達(dá)水平亦存在差異。本研究結(jié)果表明出現(xiàn)彎梗的不同切花月季品種表型性狀存在差異,原因可能與各品種間內(nèi)源激素濃度和調(diào)控各類激素的基因不同有關(guān),因此,下一步將在本研究篩選出的高盛、浪漫沙灘和艾莎3個(gè)彎梗特性較強(qiáng)的切花月季品種基礎(chǔ)上,針對(duì)調(diào)控彎梗的內(nèi)源激素及其調(diào)控基因等方面進(jìn)行深入研究。
4結(jié)論
10個(gè)切花月季品種彎梗植株表型性狀的變異較大,花梗彎曲無(wú)特定方向。氣場(chǎng)和蘇醒彎梗萼片表型變化較多;篩選出莖粗、最大花瓣厚度、花朵鮮重、花瓣數(shù)、花朵寬、花盤到第1臺(tái)葉距離、下落萼片鮮重和花梗高等8個(gè)關(guān)鍵指標(biāo)用于月季彎梗特性評(píng)價(jià);獲得高盛、浪漫沙灘和艾莎3個(gè)彎梗特性較強(qiáng)的切花月季品種,可作為研究切花月季彎梗特性的材料。
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南方農(nóng)業(yè)學(xué)報(bào)2024年9期