尚浩樂 吳娟莉 黎曼緹 朱昌威 易自力 黃紅梅
DOI:10.13925/j.cnki.gsxb.20240043
摘??? 要:【目的】掌握金柑花粉活力與柱頭可授性的變化情況,尋找最佳授粉時期,探明金柑雜交授粉的生物學(xué)特性,提高雜交成功率?!痉椒ā繉?3份金柑材料進行胚型的統(tǒng)計。選擇不同胚數(shù)材料,采用TTC染色法和聯(lián)苯胺-過氧化氫法分別對小蕾期、大蕾期、初開期和盛開期的花粉活力和柱頭可授性進行研究。對金柑進行去雄不授粉、自交授粉和雜交授粉處理,比較自交和雜交花粉管的生長情況及3種處理下的坐果率?!窘Y(jié)果】羅紋種胚數(shù)最少,適合作母本,其次是山金柑、金彈和羅浮,而長壽金柑的胚數(shù)最多;花粉活力和柱頭可授性均在4個時期呈先增后減的趨勢,并均在初開期達到了最高活性;金柑雜交花粉管和部分的自交花粉管均進入了子房;除寧波金彈和寧波羅紋CS外,其他5種金柑種質(zhì)雜交授粉比自交授粉的坐果率更高,并呈極顯著性差異;母本的柱頭活性越高,坐果率越高,種內(nèi)雜交坐果率高于種間,并呈極顯著性差異;金柑去雄不授粉下仍能得到果實,具有單性結(jié)實的特性?!窘Y(jié)論】金柑雜交中應(yīng)選擇單胚型母本材料;金柑屬于雌雄同熟,采集花粉最佳時期和授粉最佳時期均為初開期;金柑雜交親和性比自交親和性更強,有利于金柑的雜交育種。
關(guān)鍵詞:金柑;花粉活力;柱頭可授性;花粉管發(fā)育;種間雜交
中圖分類號:S666.1?????????? 文獻標(biāo)志碼:A??????????? 文章編號:1009-9980(2024)05-0875-12
收稿日期:2024-01-19??????? 接受日期:2024-03-19
作者簡介:尚浩樂,在讀碩士研究生,研究方向為金柑種質(zhì)資源利用與創(chuàng)新。E-mail:shang11244@163.com
*通信作者 Author for correspondence. E-mail:hhm7418@hunau.edu.cn
果 樹 學(xué) 報 2024,41(5): 875-886
Journal of Fruit Science
Study on the biological characteristics of cross-pollination in kumquat
SHANG Haole, WU Juanli, LI Manti, ZHU Changwei, YI Zili, HUANG Hongmei*
(Hunan Agricultural University, Changsha 410000, Hunan, China)
Abstract: 【Objective】 Pollen viability and stigma pollinability are important indicators for assessing the quality of parental gametes. Understanding the changing pattern of kumquat pollen viability and stigma pollination rate and finding the best pollination period can effectively improve the quality and success rate of hybrid breeding. Simultaneously, the growth of kumquat pollen tubes under self-pollination and heterogametic pollination conditions was studied to find out the biological characteristics and mechanism of kumquat heterogametic pollination to provide a theoretical basis for kumquat promiscuity. By studying the fruiting rate under different treatment conditions, we can provide some practical basis for kumquat promiscuity. 【Methods】 Embryo types of 23 Fortunella germplasm resources were statistically analyzed. Materials with different embryo numbers were selected to analyze pollen viability and stigma receptivity at four different stages (Small bud stage, Big bud stage, Early florescence stage and Full bloom stage) by using TTC staining and benzidine-peroxidase methods, respectively. Subsequently, three treatments, including emasculation without pollination, artificial self-pollination, and hybrid pollination were conducted. The growth of pollen tubes was compared under both artificial self-pollination and hybrid pollination. Finally, fruit setting rates after three treatments were statistically analyzed and compared. 【Results】 Fortunella japonica (Thunb) Swingle had the fewest embryos, making them suitable as female parents, followed by F. hindsii (Champ) Swingle, F. crassifolia Swingle and F. margarita (Lour.) Swingle. F. obovata Tanaka had the most embryos. Pollen viability of all tested Kumquat materials at all four stages followed the same trend of “increasing first and then decreasing” and the peak reached at the early florescence stage. The highest pollen viability was found in the Lanshanjingan (F. crassifolia Swingle), which was significantly different from other species, and Liuyangjingan CQ (F. crassifolia Swingle) had the lowest pollen viability. Stigma receptivity followed an increasing-decreasing trend at all four stages, with maximum receptivity observed at the early florescence stage. Lanshanjingan exhibited the highest pistil receptivity, while Liuyangjingan CQ showed the lowest. There were no significant differences in pistil receptivity among other kumquat varieties, and therefore they can all serve as female parents for hybridization. Analysis of pollen activity and stigma receptivity revealed that kumquats exhibited synchronous maturation of male and female reproductive organs, reaching peak values at the early florescence stage. Pollen tubes of both self-pollination and hybridization treatments began germination approximately 2 hours after pollination. Hybrid pollen tubes showed a faster elongation rate compared with self-pollination pollen tubes. After 12 to 24 hours of pollination, most hybrid pollen tubes reached the base of the style as a result of fast elongation, whereas some self-pollination pollen tubes had already ceased elongation at 2/3 of the style length. After 48 hours of pollination, all hybrid pollen tubes and a few self-pollination pollen tubes penetrated the base of the style and entered the ovary. Under the condition of emasculation without pollination, all 7 kumquat germplasm obtained fruits, and Lanshanjingan had the highest fruiting rate and showed significant difference from other kumquat germplasm. The kumquat flowers can develop into fruits without pollination, demonstrating the characteristic of parthenocarpy. Under conditions of artificial self-pollination, Lanshanjingan had the highest fruit setting rate, showing significant differences from other kumquat germplasms. Under the condition of hybridization pollination treatment, the fruiting rate of Liuyangjingan CQ was the highest, followed by Lanshanjingan. However, there was no significant difference in fruit set rate between these two kumquat varieties, but there was a significant difference between them and among the other 5 kumquat varieties. Except for Ningboluowen CS [F. japonica (Thunb) Swingle], the fruit setting rates of the other 6 kumquat from cross-pollination treatments were higher than those from artificial self-pollination; among the 7 kumquat varieties, only Ningbojindan (F. crassifolia Swingle) showed no significant difference in fruit setting rates under both artificial self-pollination and cross-pollination treatments. The difference in fruit set rate indicated that the cross-compatibility of kumquats was stronger than self-compatibility, which was more conducive to the hybrid breeding of kumquats. With the same hybrid paternal parent, the higher the stigma pollinability of the female parent, the higher the fruit setting rate. The fruit setting rate of intraspecific hybridization was higher than that of interspecific hybridization, and there was a significant difference. 【Conclusion】 The female parent with single embryo should be selected in kumquat hybridization; kumquat belonged to hermaphroditism, and the best time for pollen collection and pollination was at the early florescence stage; kumquat cross-compatibility was stronger than self-compatibility, which was beneficial to kumquat cross breeding.
Key words: Kumquat; Pollen viability; Stigma pollinability; Pollen tube development; Interspecific hybridization
金柑屬(Fortunella Swingle,1915)植物,別名金橘等,原產(chǎn)中國,為蕓香科(Rutacease)柑橘亞科(Aurantioideae)柑橘族(Citreae)柑橘亞族(Citrinae)的一類植物。主要包括金豆(F. hindsii Swingle)、羅紋(F. japonica Swingle)、長葉金柑(F. polyandra Tanaka)、羅浮(F. margarita Swingle)4個種和長壽金柑(F. obovata Tanaka)、金彈(F. crassifolia Swingle)2個雜交種[1],在廣西融安、陽朔、湖南瀏陽和藍山等地廣泛種植[2-3],食用價值和藥用價值極高[4-9],國內(nèi)外市場競爭力強[10-13],具有廣闊的發(fā)展前景。
目前,果樹育種主要采用雜交育種和芽變育種兩個途徑。據(jù)不完全統(tǒng)計,在中國的蘋果、桃及梨等11種果樹中,約有63.9%的新品種是通過雜交育種獲得的[14]。然而,柑橘類植物的育種仍以芽變育種為主,通過雜種選育得到的新品種僅占6.3%[15]。雜交育種可以從父母本獲得優(yōu)良性狀,能夠滿足人們對培育果樹新品種的需求,因此傳統(tǒng)的人工雜交育種技術(shù)在培育新品種方面仍具有重要地位[16]。
在植物的雜交育種中,花粉是遺傳物質(zhì)傳遞的中間媒介[17],掌握花粉的生物學(xué)特性可以有效提高授粉成功率,對選擇合適的雜交親本意義重大[18],而柱頭的可授性則直接影響到受精能否繼續(xù)進行[19]。因此,研究花粉柱頭活性,選擇優(yōu)良的雜交親本,可以有效提高結(jié)實率,有助于培育雜交新品種[20]。近年來,學(xué)者們對花粉活力和柱頭可授性等的研究越發(fā)深入,為雜交育種提供了堅實的理論支持;Liu等[21]以擬南芥為研究對象,揭示了花粉-柱頭保守的識別機制,這對克服遠緣雜交障礙、獲得優(yōu)良性狀的品種具有重要意義;朱江華等[22]通過對比研究6個藍莓品種的花粉量、花粉活力及花粉萌發(fā)率等花粉特性,發(fā)現(xiàn)花粉量大、萌發(fā)率高、花粉活力好的品種在授粉過程中表現(xiàn)出特殊的優(yōu)異性。
筆者在本研究中通過比較不同時期的花粉活力與柱頭可授性,確定最佳授粉時期。對金柑進行去雄不授粉、自交授粉和雜交授粉三種處理并比較自交授粉和雜交授粉的花粉管的生長情況及三種處理下的坐果率,初步探明金柑雜交授粉的生物學(xué)特性,為金柑雜交育種提供一定的理論依據(jù)。
1 材料和方法
1.1 試驗材料
試驗材料采自湖南農(nóng)業(yè)大學(xué)金山金柑資源圃,其中,金彈種15份材料,羅紋種3份材料,羅浮種2份材料,山金柑種2份材料和長壽金柑種1份材料,共計23份材料(表1)。
1.2 試驗方法
試驗于2020年6—8月在金山基地進行。并將金柑開花動態(tài)分為以下4個時期[23],每個時期對應(yīng)的花蕾形態(tài)如圖1。
1.2.1 自然授粉果實胚數(shù)統(tǒng)計 在自然授粉結(jié)實條件下,從23份種質(zhì)資源成熟果實中取出種子,去除種皮,將子葉分開觀察胚數(shù),每種材料至少5個果實,計算種子數(shù)與胚數(shù)的平均值。按照胚數(shù)從少到多分別選擇羅紋種的寧波羅紋、寧波羅紋CQ和寧波羅紋CS,金彈種的寧波金彈、寧波金彈CQ、瀏陽金柑CQ、溫州金彈CS、藍山金柑和瀏陽金柑,羅浮種的牛奶金柑用于后續(xù)試驗。
1.2.2 花粉活力的測定 參照0.5%TTC染色法[24]并略作修改,分別在金柑花蕾的小蕾期、大蕾期、初開期和盛開期4個時期,各采集10個花蕾樣本。取花藥置于載玻片上,滴加1~2滴0.5%TTC溶液(2,3,5-uiphenyl tetrazolium chloride),置于濕潤的培養(yǎng)皿中,室溫下靜置30~45 min,在顯微鏡視野中隨機選擇5個不同的位置,統(tǒng)計總花粉粒數(shù)和變色花粉粒數(shù)(包括紅色和粉色)。按照公式1計算花粉活力。
[花粉活力/%=變色花粉數(shù)觀察花粉總數(shù)×100]。????????? (1)
1.2.3 柱頭活性的測定 參照聯(lián)苯胺-過氧化氫法[25],分別在金柑花蕾的小蕾期、大蕾期、初開期和盛開期4個時期,各采集10個花蕾樣本,單獨取柱頭于卡諾式固定液中固定,取出柱頭,浸入滴有聯(lián)苯胺-過氧化氫溶液的凹面載玻片中,室溫靜置10 min后在顯微鏡下觀察柱頭周圍的氣泡產(chǎn)生及顏色變化情況。柱頭可授性檢測標(biāo)準(zhǔn)參照焦雪輝等[24]的方法(表2)。
1.2.4 花粉的采集與授粉 基于花粉活力與柱頭可授性的試驗結(jié)果,選定寧波金彈、瀏陽金柑CQ、寧波羅紋、寧波羅紋CQ、藍山金柑、寧波羅紋CS、牛奶金柑及溫州金彈CS,共計8種材料為雜交親本。自交及雜交組合情況詳見表3。
每天早晨07:00—08:30采集父本花蕾接近初開期之前的雄蕊。取花藥并在干燥器中干燥至其花粉散落,4 ℃冰箱中保存?zhèn)溆谩?/p>
每天上午07:00—09:00進行授粉。
(1)去雄不授粉:去除母本花蕾接近初開期之前的雄蕊,套上硫酸紙袋并記錄去雄時間,3 d后取下硫酸袋,10 d后統(tǒng)計坐果率。
(2)自交授粉:授粉前1 d下午18:00,去除目標(biāo)材料花瓣和雄蕊,套上硫酸袋。第2天早上07:00,用毛刷進行人工授粉。授粉時,用毛筆刷輕輕地將花粉刷在柱頭上,直到柱頭表面明顯觀察到黃色花粉,確保授粉成功。每朵花連續(xù)3 d在相同時間授粉3次,從第3天授粉完成后開始計算,3 d后去袋,10 d后統(tǒng)計坐果率。
(3)雜交授粉:按照表3的雜交組合,將處理好的父本花粉涂抹在選定的母本柱頭上,其他操作與自交授粉相同。
1.2.5 花粉管生長觀察 在大蕾期,按照表3所示的自交與雜交組合分別進行授粉處理,授粉方法與1.2.4節(jié)相同,在授粉結(jié)束后的2、4、8、12、24、48 h分別取10個花柱從子房基部切除,于卡諾式固定液中固定12~24 h。之后分別經(jīng)過70%乙醇、50%乙醇、30%乙醇及蒸餾水沖洗后,放于2 mol·L-1的NaOH溶液中軟化8~12 h。最后在0.1%水溶性苯胺藍中染色12~24 h,用解剖刀切開柱頭并壓片,在熒光顯微鏡下觀察花粉管生長情況并拍照。
1.2.6 坐果率的統(tǒng)計 統(tǒng)計各種處理下的授粉花朵數(shù),并在授粉完成后的第10天統(tǒng)計坐果數(shù),按照公式2計算坐果率。
[坐果率/%=坐果數(shù)授粉花朵數(shù)×100]。?????????????? (2)
1.2.7 數(shù)據(jù)處理與分析 本試驗數(shù)據(jù)通過Excel進行記錄、統(tǒng)計,采用SPSS 26.0進行方差分析,用Origin 2018進行圖表繪制。
2 結(jié)果與分析
2.1 金柑種質(zhì)資源胚數(shù)分析
23份種質(zhì)的種子數(shù)及胚數(shù)統(tǒng)計結(jié)果如表4,羅紋種胚數(shù)均為1個,為單胚型種質(zhì),最適合做母本;其次為山金柑、金彈和羅浮,胚數(shù)最多的為長壽金柑。
2.2 金柑不同時期花粉活力比較
不同金柑花蕾在4個時期的花粉活力均呈現(xiàn)先升后降的趨勢,并在初開期達到了最強水平(圖2)。相同時期下,藍山金柑的花粉活力最強,與除溫州金彈CS以外的其他金柑種質(zhì)存在顯著差異(p<0.05)。在大蕾期,寧波金彈、溫州金彈CS和瀏陽金柑CQ間不存在顯著性差異(p>0.05),且該3種金柑與藍山金柑、寧波羅紋CQ和瀏陽金柑呈顯著性差異(p<0.05)。
雜交中為避免串粉現(xiàn)象,選擇接近初開期的大蕾期為采集花粉的時期。在大蕾期,金柑花粉活力從高到低分別為藍山金柑>瀏陽金柑CQ>溫州金彈CS>寧波金彈>牛奶金柑>寧波羅紋CQ>瀏陽金柑。選擇花粉活力較強的種質(zhì)作為雜交授粉的父本,即藍山金柑、瀏陽金柑CQ、溫州金彈CS。
2.3 金柑不同時期柱頭可授性比較
不同金柑花蕾在4個時期的柱頭可授性均呈現(xiàn)先增強后減弱的趨勢,并在初開期達到了最高水平(圖3)。在大蕾期時,藍山金柑的柱頭可授性高于其他金柑并存在顯著性差異(p<0.05),除藍山金柑外的其他7種金柑間不存在顯著性差異(p>0.05)。
柱頭可授性的強弱會直接影響花粉在柱頭上的萌發(fā)概率,柱頭可授性越強,接受花粉的能力越強,從而更容易完成受精過程。通過對金柑4個時期柱頭活性的檢測,發(fā)現(xiàn)金柑在初開期普遍表現(xiàn)出最強的柱頭可授性。所測材料均適用作人工雜交授粉的母本材料。然而,為了避免串粉現(xiàn)象,應(yīng)在接近初開期之前去除雄蕊并授粉。
2.4 金柑自交與雜交花粉管生長比較
通過對自交和雜交的花粉管進行觀察和比較,發(fā)現(xiàn)所有的自交和雜交花粉管在授粉后2 h左右開始萌發(fā)(圖4-A、B);授粉4~8 h時,雜交花粉管的生長長度達到了花柱總長的1/3~1/2,而自交花粉管生長速度相對較慢;授粉12~24 h時,大多數(shù)雜交花粉管已經(jīng)延伸到了花柱基部(圖4-C),一些自交花粉管在花柱總長的2/3處停止了伸長(圖4-D)。授粉48 h后,所有雜交花粉管和部分自交花粉管穿過花柱基部,進入子房內(nèi)部(圖4-E、F),寧波金彈、寧波羅紋CS及牛奶金柑等自交組合中,花粉管生長緩慢甚至停止。上述結(jié)果表明金柑種質(zhì)具有完全雜交親和性,且存在部分自交不親和現(xiàn)象。
2.5 不同處理條件下坐果率的比較
在去雄不授粉條件下,7份金柑種質(zhì)均得到了果實,說明金柑子房在未授粉的情況下也能發(fā)育成為果實,具有單性結(jié)實的特性;在人工自交授粉處理條件下,藍山金柑的坐果率最高,并與其他金柑種質(zhì)間呈現(xiàn)出顯著性差異;在雜交授粉處理條件下,瀏陽金柑CQ的坐果率最高,其次為藍山金柑,該2種金柑種質(zhì)間不存在顯著性差異,但與其他5種金柑種質(zhì)存在顯著性差異(圖5)。在人工自交授粉、去雄不授粉和雜交授粉三種處理條件下,藍山金柑得到了較高的坐果率,與藍山金柑花粉活性和柱頭可授性均最高有關(guān)。
同一母本材料在人工自交授粉處理及雜交授粉處理下的坐果率差異性分析如表5,發(fā)現(xiàn)除寧波金彈和寧波羅紋CS外,其他5種金柑種質(zhì)在兩種不同處理條件下的坐果率呈現(xiàn)出極顯著性差異,且雜交授粉比自交授粉的坐果率更高,說明金柑種質(zhì)的雜交親和性比自交親和性更強,有利于金柑的雜交育種。上述結(jié)果與2.4花粉管生長的試驗結(jié)果相同。
2.6 金柑雜交授粉的坐果率差異性分析
在雜交授粉處理條件下,同一父本不同母本間坐果率的差異情況如表6所示。父本材料為溫州金彈CS時,母本材料為寧波金彈與寧波羅紋坐果率呈現(xiàn)出極顯著性差異,柱頭活性高的寧波羅紋坐果率更高;母本材料寧波金彈與寧波羅紋CQ坐果率呈現(xiàn)出極顯著性差異;寧波羅紋和寧波羅紋CQ間坐果率差異并不顯著,但寧波羅紋的柱頭活性更強,坐果率更高;父本材料為藍山金柑時,母本材料為寧波羅紋CS與瀏陽金柑CQ的坐果率存在極顯著性差異,盡管寧波羅紋CS的柱頭活性更高,但坐果率卻更低,說明種間雜交親和性更差;父本材料為瀏陽金柑CQ時,母本材料牛奶金柑和藍山金柑的坐果率間存在極顯著性差異,藍山金柑柱頭活性更強,且種內(nèi)雜交親和性更強,因而其坐果率更高。綜上所述,柱頭活性越高,坐果率越高,且種內(nèi)雜交的坐果率高于種間雜交。因此在選擇雜交親本時,應(yīng)考慮柱頭活性和親本親緣關(guān)系等因素。
3 討 論
了解植物授粉生物學(xué)特性對植物雜交育種具有重要的指導(dǎo)意義[20,25-27]。對于柑橘類植物的雜交育種來說,經(jīng)常由于珠心胚的干擾而不能獲得雜種,其原因是珠心胚與合子胚之間存在競爭關(guān)系,導(dǎo)致合子胚發(fā)育失敗[28]。但當(dāng)以單胚為母本時可以提高柑橘類植物的雜種率,因此筆者在本研究中比較了不同金柑種質(zhì)的胚數(shù),以篩選出單胚種質(zhì),作為雜交的母本材料。筆者在本研究中發(fā)現(xiàn)羅紋種為單胚種質(zhì),適合作為雜交母本。
柱頭可授性和花粉活性也是影響雜交成功率的重要因素。通過比較不同時期金柑花蕾的柱頭可授性和花粉活性,結(jié)果顯示二者的活性趨勢相似,且均在初開期達到最強,屬于雌雄同熟植物,并未發(fā)現(xiàn)花粉敗育情況[29]。試驗結(jié)果與百香果和櫻桃相同[30-31]。但是在雜交實踐中,為了避免串粉,通常在初開期之前的大蕾期進行花粉采集和去雄處理。
被子植物受精過程主要包括花粉與柱頭的識別、花粉管在花柱間的伸長,以及雌雄配子間的融合三個階段,任意階段無法識別均會導(dǎo)致雜交失敗[32]。在授粉后2 h內(nèi)為花粉與柱頭識別的重要時間段[33],筆者在本研究中發(fā)現(xiàn),金柑在授粉處理2 h后,所有自交和雜交均完成了花粉與柱頭的識別,花粉管開始在花柱內(nèi)生長。在授粉48 h后,所有雜交花粉管和部分自交花粉管穿過花柱基部進入子房。在花粉管的行為上表現(xiàn)出了較好的雜交親和性,該現(xiàn)象在多種植物中被發(fā)現(xiàn)[34-35]。在本研究中,自交花粉管在花柱內(nèi)停滯生長,表現(xiàn)出自交不親和的現(xiàn)象,同樣被雷翠云[36]發(fā)現(xiàn)。在花粉管生長階段導(dǎo)致果樹自交不親和的因素有很多[37],目前對金柑中花粉管生長停滯現(xiàn)象的原因尚未見報道。在柑橘類植物中,被證實S-RNase基因的表達調(diào)控[38]和信號傳導(dǎo)[39]均會對花粉管的發(fā)育產(chǎn)生影響。其他原因如花粉管的導(dǎo)向生長[40]、微絲、微管骨架[41]等也會導(dǎo)致花粉管的生長停滯。
坐果率能直接反映出種質(zhì)間親和性的強弱,是研究種質(zhì)間雜交及自交親和性的重要指標(biāo)之一[42-44]。筆者在本研究中發(fā)現(xiàn),大部分金柑雜交的坐果率高于自交,并呈現(xiàn)極顯著差異,該現(xiàn)象的原因可能是金柑具有雜交親和性以及部分自交不親和性,并且與花粉管的生長觀察試驗結(jié)果相同。該現(xiàn)象已在植物中被發(fā)現(xiàn),如梨樹[45]、甜菊[46]、棗[47]等。并且筆者在本研究中還發(fā)現(xiàn),對金柑進行去雄不授粉處理后,子房仍可以發(fā)育成果實,表明金柑具有單性結(jié)實的特性,與雷翠云[36]的研究結(jié)果相同。
植物柱頭的可授性強弱可以直接影響坐果率的高低,也是決定雜交育種工作成功與否的重要指標(biāo)之一[48-49]。筆者在本研究中發(fā)現(xiàn)柱頭活性越強,坐果率越高,且種內(nèi)雜交的坐果率高于種間雜交。相關(guān)研究也表明,柱頭可授性與坐果率間存在顯著正相關(guān)[26,50]。因此在選擇雜交親本時,應(yīng)考慮柱頭活性和親本親緣關(guān)系等因素。以上對金柑雜交的生物學(xué)特性研究,為金柑雜交條件的確定以及雜交親本的選擇提供了一定的參考依據(jù)。
4 結(jié) 論
金柑雜交中應(yīng)選擇單胚型母本材料;金柑屬于雌雄同熟果樹,采集花粉最佳時期和授粉最佳時期均為初開期;金柑雜交親和性比自交親和性更強,有利于金柑的雜交育種。
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