趙永昌 柴紅梅 陳衛(wèi)民 趙子悅 孔令艦

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羊肚菌子實(shí)體發(fā)育生物學(xué)（下）——生物學(xué)和非生物學(xué)因子對(duì)菌絲培養(yǎng)和子實(shí)體形成的影響

趙永昌 柴紅梅 陳衛(wèi)民 趙子悅 孔令艦

（云南省農(nóng)業(yè)科學(xué)院生物技術(shù)與種質(zhì)資源研究所，云南 昆明 650221）

3 羊肚菌發(fā)育生物學(xué)

羊肚菌的生活史是研究的難點(diǎn)，目前有提出兩種生活史[114, 115]，都認(rèn)為形成菌核和分生孢子是關(guān)鍵環(huán)節(jié)。儲(chǔ)存營(yíng)養(yǎng)的菌核在子囊果發(fā)育過程中作用較大，既可以萌發(fā)為生殖菌絲，又可以直接產(chǎn)生子囊果。羊肚菌的菌核為假菌核，不同的可栽培出菇的菌株產(chǎn)菌核能力差別較大，甚至有菌種階段完全不產(chǎn)菌核的菌株也能很好地出菇；另一方面尚無菌核萌發(fā)成原基的直接證據(jù)。分生孢子的爭(zhēng)議則較大，主要是從自然基質(zhì)得到的分生孢子不能萌發(fā)，且羊肚菌產(chǎn)量與分生孢子產(chǎn)生量沒有直接相關(guān)性。使用覆膜技術(shù)種植羊肚菌有的分生孢子非常少但出菇不受影響，用圓葉楊種植也幾乎未發(fā)現(xiàn)有分生孢子形成。因此，羊肚菌的發(fā)育生物學(xué)基礎(chǔ)研究可能是解決羊肚菌穩(wěn)產(chǎn)、育種的關(guān)鍵，而實(shí)現(xiàn)室內(nèi)穩(wěn)定出菇則是研究的基礎(chǔ)。解決羊肚菌培養(yǎng)過程中遺傳的多變性是重中之重。

3.1 子囊孢子形成與萌發(fā)

子囊孢子的形成與萌發(fā)是羊肚菌發(fā)育的關(guān)鍵問題，羊肚菌子囊孢子具有多核現(xiàn)象，有的多達(dá)數(shù)十個(gè)核，這些核是同核體還是異核體尚不清楚。雖然有不少學(xué)者試圖對(duì)羊肚菌子囊孢子的形成進(jìn)行深入研究，但是由于未實(shí)現(xiàn)室內(nèi)栽培，難以找到不同發(fā)育時(shí)期的子囊孢子個(gè)體，導(dǎo)致研究困難。陳立佼借助梯棱羊肚菌的大田栽培，通過細(xì)胞核染色技術(shù)對(duì)子囊孢子的形成進(jìn)行研究，總結(jié)出子囊孢子形成的過程模式[116]，說明羊肚菌子囊孢子的多核是孢子形成后的有絲分裂導(dǎo)致的，多核子囊孢子應(yīng)是同核體，這方面的結(jié)果得到交配型基因鑒定證明。Chai等證明-20和子囊孢子群都是同核體，具有1-1和1-2交配型基因子囊孢子的比例與1?1無偏差，所以M-20和是異宗結(jié)合的[117]。有研究認(rèn)為栽培條件下子囊果形成孢子比較晚，自然生長(zhǎng)的子囊果傾斜或倒伏狀態(tài)時(shí)才會(huì)形成孢子，此時(shí)子囊內(nèi)除孢子外基本呈透明狀態(tài)，不存在其他細(xì)胞器或大分子。也有不形成孢子的共生類型的羊肚菌菌株[118]。

在室內(nèi)條件下，羊肚菌的子囊孢子較易萌發(fā)，在濕度適宜時(shí)3小時(shí)就可見到芽管，因子囊孢子的多核性其可能從1-5端長(zhǎng)出芽管萌發(fā)；在野外，影響子囊孢子萌發(fā)的因素很多，溫度、濕度、營(yíng)養(yǎng)條件都可能影響。研究表明，子囊孢子萌發(fā)對(duì)溫度比較敏感，室內(nèi)溫度低至2 ℃都能萌發(fā)，但將孢子包埋在玻璃紙里放置在出菇點(diǎn)時(shí)，土壤溫度超過10 ℃都不萌發(fā)。當(dāng)土壤持續(xù)升溫到15 ℃時(shí)會(huì)抑制子囊孢子的萌發(fā)，地表的孢子一年后將失去活力。將經(jīng)滅菌的黑麥埋入地中，土壤溫度保持在10 ℃以下時(shí)，無論孢子還是菌核都容易在其上定植[119]。一定濃度范圍的MgSO4、KNO3、CaCl2、NaNO3對(duì)孢子萌發(fā)和菌絲生長(zhǎng)有促進(jìn)作用[120]。子囊孢子萌發(fā)的菌絲一般仍為多核體細(xì)胞，同一子囊孢子萌發(fā)得到的初生菌絲不發(fā)生融合，也未觀察到不同子囊孢子萌發(fā)的菌絲之間發(fā)生融合[118, 121]。有研究表明，單孢菌株配對(duì)后能形成異核體，這與菌柄分離的菌株為異核體類似，間接表明其為異宗結(jié)合[114, 122, 123]。

3.2 多變的培養(yǎng)形態(tài)

羊肚菌的培養(yǎng)形態(tài)變異之大是目前大型真菌中少有的，相同培養(yǎng)條件下不同種或同種不同菌株在菌核形成的時(shí)間、形態(tài)、多少和大小等方面均有不同，在不同的培養(yǎng)條件下同一菌株菌核的產(chǎn)生和形態(tài)也不同，培養(yǎng)過程中的變化不僅類型多且重復(fù)性較差。雖然培養(yǎng)基組成、生長(zhǎng)條件顯著影響羊肚菌生長(zhǎng)速度[124～129]，但引起這些變化的原因是遺傳性的還是純粹生理性的尚不清楚。羊肚菌菌絲存在多核現(xiàn)象，這種多核仍舊是同核體。同核體菌株培養(yǎng)過程中變化穩(wěn)定，不存在由于核丟失而產(chǎn)生變化。

陳立佼等從單孢群體出發(fā)以產(chǎn)菌核能力為定性特征研究了培養(yǎng)特性變化[130]，單孢菌株按培養(yǎng)特征可分為9類（表3），同等條件下每一菌株的培養(yǎng)特性保持穩(wěn)定，在綜合馬鈴薯葡萄糖培養(yǎng)基（CPDA）、葡萄糖硝酸鈉瓊脂培養(yǎng)基（GN）、酵母膏胨葡萄糖瓊脂培養(yǎng)基（YPD）進(jìn)行轉(zhuǎn)接培養(yǎng)時(shí)，可成功地將產(chǎn)菌核菌株轉(zhuǎn)化為不產(chǎn)菌核菌株。同一子實(shí)體及不同子實(shí)體產(chǎn)菌核單孢菌株產(chǎn)核數(shù)量及分布，變化很大，對(duì)峙培養(yǎng)后單孢之間性狀會(huì)發(fā)生較大變化，包括菌核形態(tài)、菌絲形態(tài)、生長(zhǎng)勢(shì)，特別是產(chǎn)菌核能力會(huì)消失和發(fā)生轉(zhuǎn)移：①同一子實(shí)體單孢菌株之間存在拮抗作用，對(duì)峙培養(yǎng)時(shí)相互接觸部位會(huì)形成交合線，出現(xiàn)濃密的生長(zhǎng)區(qū)，將不產(chǎn)菌核的單孢菌株與同一子實(shí)體及不同子實(shí)體產(chǎn)菌核單孢菌株進(jìn)行對(duì)峙培養(yǎng)，即使來自相同子實(shí)體的單孢菌株之間也會(huì)出現(xiàn)拮抗作用。拮抗線兩側(cè)的菌絲和親本菌株相比形態(tài)和產(chǎn)核情況有很大變化，在各對(duì)峙組合中不同親本的組合拮抗線情況不同。②在對(duì)峙組合中由于拮抗作用，兩親本的生長(zhǎng)勢(shì)不同，有拮抗的組合中產(chǎn)菌核菌株長(zhǎng)勢(shì)明顯占優(yōu)，只是有的長(zhǎng)勢(shì)處于弱勢(shì)，可能是由于產(chǎn)菌核菌株菌絲輻射狀快速生長(zhǎng)，使其在平板上能迅速占據(jù)更多培養(yǎng)資源。③產(chǎn)菌核水平變化，拮抗作用對(duì)產(chǎn)菌核能力影響較大，有產(chǎn)核能力減弱的，有完全失去產(chǎn)菌核能力的，也有產(chǎn)菌核能力互換的（在組合中一個(gè)菌株將產(chǎn)菌核能力“轉(zhuǎn)染”，原來產(chǎn)菌核的菌株仍然產(chǎn)菌核，不產(chǎn)菌核的菌株能產(chǎn)菌核），可推測(cè)在對(duì)峙培養(yǎng)過程中與菌核形成相關(guān)的遺傳因子可在細(xì)胞間遷移，或是菌核產(chǎn)生與某種分泌物有關(guān)。該物質(zhì)有一定的趨向性，各菌株對(duì)其吸引能力的強(qiáng)弱決定了它的分布，從而影響了菌核的產(chǎn)生和分布。

表3 基于YPD上的培養(yǎng)特征的羊肚菌單孢菌株分組[130]

3.3 菌核形成與變化

3.3.1 純培養(yǎng)條件下菌核變化

菌核是一種由菌絲組織化形成的具有抵御逆境作用的致密菌絲結(jié)構(gòu)，逆境是菌核產(chǎn)生的重要基礎(chǔ)與條件；然而羊肚菌在營(yíng)養(yǎng)豐富的環(huán)境中也能產(chǎn)生菌核。說明營(yíng)養(yǎng)因素既可以刺激也可能抑制羊肚菌菌核的形成。盡管羊肚菌栽培取得了進(jìn)展，但仍具有挑戰(zhàn)性，因?yàn)榧词拱凑諏＠膊荒芡耆耘喑龉?。多?shù)研究認(rèn)為，要獲得栽培成功，必須獲得菌核，不少人曾經(jīng)將精力集中在菌核產(chǎn)生研究上。對(duì)于羊肚菌菌核的形態(tài)和形成過程有不同看法，Alvarado-Castillo等[131]的分類比較合理，包括七個(gè)階段：①菌絲生長(zhǎng)，②初級(jí)菌絲生長(zhǎng)和分枝，③次級(jí)菌絲纏繞，④大量菌絲聚集，⑤菌絲聚合生長(zhǎng)，⑥菌核形成，⑦菌核生長(zhǎng)與成熟，成熟伴隨厚垣孢子和分生孢子形成。

影響菌核形成的因素較多，包括環(huán)境因子和營(yíng)養(yǎng)因子[71, 132]，營(yíng)養(yǎng)因子涉及種類和濃度。在不同蔗糖濃度的培養(yǎng)基上，菌絲生長(zhǎng)速度、菌核顏色差別較大，且形成菌核的菌絲體外表會(huì)有一些具備儲(chǔ)蓄能力的氣泡[133, 134]。氮源的影響高于碳源[135]，在含核糖、半乳糖、山梨糖和甘露醇的培養(yǎng)基上較難形成菌核，而NaNO3和酵母提取物有利于分散型菌核形成[136, 137]。復(fù)合營(yíng)養(yǎng)物如玉米、小麥、燕麥和黑麥等，對(duì)菌核的形成有促進(jìn)作用[138]。而饑餓可以誘導(dǎo)和增加假菌核的形成，假菌核中丙二醛含量增加與脂肪的過氧化和氧化壓力有關(guān)[139]。影響菌核形成的非營(yíng)養(yǎng)因素包括光照、溫度、有機(jī)酸、環(huán)境腐敗產(chǎn)物的積累、酚類化合物、多酚氧化酶活性、機(jī)械屏障、疏基修飾因子，以及滲透壓等，能增加或減弱氧化應(yīng)激的生長(zhǎng)因子可能都有促進(jìn)或抑制菌核分化的作用。

3.3.2 菌核生理變化

木質(zhì)素降解酶是大型真菌最重要的酶之一，其參與人工栽培的子實(shí)體形成。在羊肚菌生活史中，菌核是羊肚菌從菌絲到子實(shí)體的中間態(tài)。研究表明，在不同培養(yǎng)基上，羊肚菌菌核形成過程中木質(zhì)素降解酶的活性有較大變化，不同培養(yǎng)基上都產(chǎn)生漆酶（LAC）、木質(zhì)素過氧化物酶（LiP）和錳過氧化物酶（MnP）等木質(zhì)素降解酶，在麥粒培養(yǎng)基上LAC活性最高，而在谷殼培養(yǎng)基上則是MnP和LiP最高；在土壤復(fù)合培養(yǎng)基上，從菌核形成到成熟都能產(chǎn)生漆酶，都能測(cè)到漆酶活性[140, 141]。由于基質(zhì)對(duì)菌核形成及相關(guān)酶分泌有誘導(dǎo)作用，所以羊肚菌栽培過程中栽培種配方的選擇對(duì)菌核形成及后期胞外酶的表達(dá)有決定性作用。

3.3.3 菌核形成過程中基因表達(dá)變化

羊肚菌菌核形成能力的差異，表面上受環(huán)境因素影響，實(shí)際上是由菌株本身的遺傳多樣性決定的。Chen等[142]利用mRNA差異顯示技術(shù)對(duì)同一子實(shí)體分離的在相同培養(yǎng)條件下得到的產(chǎn)菌核和不產(chǎn)菌核的羊肚菌單孢進(jìn)行研究，得到了陽性的假設(shè)蛋白及未知的差異片段58個(gè)，其中26條片段為已知蛋白功能片段、9條與已知物種的假設(shè)蛋白編碼片段相似，23條為未找到假設(shè)蛋白片段。這些片段對(duì)應(yīng)的基因可能控制著羊肚菌產(chǎn)菌核與否或能力高低，為在遺傳和基因調(diào)控水平研究羊肚菌產(chǎn)菌核的機(jī)理提供了重要的遺傳信息。在確定功能的基因中，氨基丁酸通透酶片段、外膜家族蛋白相關(guān)蛋白片段和人類角蛋白結(jié)合蛋白片段都與羊肚菌逆境應(yīng)激和結(jié)構(gòu)適應(yīng)性變化有關(guān)，即得到的差異片段與植物中的脂質(zhì)代謝、氮代謝、逆境應(yīng)激及細(xì)胞周期調(diào)控有關(guān)。這些片段可能是羊肚菌的氧化應(yīng)激反應(yīng)的上游誘導(dǎo)因子或下游產(chǎn)物，需要得到片段全長(zhǎng)并進(jìn)行更深入的功能研究才能確定。表明羊肚菌菌核形成能力可能更多地與菌株對(duì)外界環(huán)境適應(yīng)性的遺傳多樣性有關(guān)。

3.3.4 菌核與子實(shí)體形成能力之間的關(guān)系

雖然最早的研究認(rèn)為，菌核是羊肚菌生活史中最重要的階段[9]，易出菇的火燒地在兩年內(nèi)觀測(cè)到的羊肚菌菌核比非火燒地多得多[143]。但對(duì)羊肚菌的生活環(huán)境調(diào)查研究發(fā)現(xiàn)，菌核并不是所有種的羊肚菌生活史的必須階段，不同種在生活史的進(jìn)化上有區(qū)別[71]。目前實(shí)驗(yàn)證明，菌核的有無與出菇與否及產(chǎn)量高低之間沒有必然的聯(lián)系，有菌核產(chǎn)生的菌株不一定出菇，很少形成菌核的菌株同樣也可以出菇[144]。雖然在栽培實(shí)踐中，多數(shù)種植者認(rèn)為需要栽培種長(zhǎng)滿菌袋（瓶）并形成菌核后才可以播種，但不少菌絲長(zhǎng)滿后就播種的出菇時(shí)間縮短，且產(chǎn)量穩(wěn)定。

3.4 分生孢子

羊肚菌栽培過程中會(huì)形成分生孢子，Ower[9]也有提及。分生孢子在目前假設(shè)的羊肚菌生活史中是重要的階段。研究表明，羊肚菌分生孢子多數(shù)為單核[145]，且產(chǎn)生與否與出菇與否無直接關(guān)系[144]。但也有認(rèn)為分生孢子類似于植物花粉的作用，對(duì)出菇有重要的意義[146]。

圖2 羊肚菌產(chǎn)生分生孢子（左）與不產(chǎn)生分生孢子（右）

明確羊肚菌分生孢子的生物學(xué)功能，首先須解決無菌培養(yǎng)條件下分生孢子的形成和分生孢子的萌發(fā)兩個(gè)難題。分生孢子可能是一種休眠體，其萌發(fā)需要特定的環(huán)境條件，目前尚無在栽培過程中觀察到分生孢子萌發(fā)的報(bào)道。根據(jù)我們研究，在滅菌基質(zhì)上生長(zhǎng)的羊肚菌可以形成分生孢子（圖2）：①分生孢子形成與器皿無關(guān)，不同的器皿培養(yǎng)都能形成分生孢子；②分生孢子形成與接種量密切相關(guān)，當(dāng)接種量大時(shí)不形成分生孢子；③分生孢子形成與溫度相關(guān)，溫度超過24 ℃，形成的分生孢子較少，而10～24 ℃間的差別不明顯，但形成時(shí)間有差異；④在最優(yōu)條件下，不同菌株分生孢子的產(chǎn)生量差別較大，同一物種分生孢子形成數(shù)量多孢菌株>單孢菌株>組織分離菌株；⑤分生孢子形成與土壤基質(zhì)的顆粒度密切相關(guān)，顆粒度小分生孢子數(shù)量少。

A：分散型原基；B：叢生型原基；C：分散性菌核周邊的原基；D：大菌核周邊的原基。

3.5 原基形成分化與子囊果形成

Ower[9]首次較為完整地描述了羊肚菌子實(shí)體，之后不斷有人對(duì)羊肚菌發(fā)育過程進(jìn)行描述[69,70,144]，基本特征是在基質(zhì)表面上從一個(gè)點(diǎn)逐步形成菌絲團(tuán)，菌絲團(tuán)上下組織化分化出菌蓋和菌柄，菌柄往下生長(zhǎng)吸收營(yíng)養(yǎng)，菌蓋往上發(fā)育逐步成熟。與其他菇類原基下面是密集的菌絲不同，大田中經(jīng)常觀察到羊肚菌原基是由一個(gè)小點(diǎn)形成的，羊肚菌子實(shí)體發(fā)育的營(yíng)養(yǎng)是如何供給輸送的尚不清楚。我們研究發(fā)現(xiàn)光溫濕可控條件下原基可以形成（圖3）：①單一交配型的菌株都不能形成原基，不同物種多孢分離菌株和組織分離菌株在原基形成的形態(tài)和數(shù)量上有差異，但無規(guī)律；②原基形成的數(shù)量與分生孢子形成相關(guān)，沒有分生孢子（包括接種量大、24 ℃培養(yǎng)）的都未形成原基；③雖然很難看到原基由菌核分化而來，但菌核多的原基數(shù)量多且密。雖然是初步結(jié)果，且在原基的分化和幼菇發(fā)育控制上尚需進(jìn)一步研究，但結(jié)合大田的種植結(jié)果可以為羊肚菌優(yōu)良菌株鑒定體系的建立奠定基礎(chǔ)。

圖4 羊肚菌栽培兩種不同的出菇方式

3.6 出菇方式與富營(yíng)養(yǎng)栽培

大田栽培羊肚菌出菇方式有見原基和不見原基兩種（圖4），有以下特點(diǎn)：①與菌株特性有關(guān)，多數(shù)菌株是見原基出菇，有的則易不見原基出菇，也有兩種出菇形式都常見；②與土壤的酸堿度、顆粒大小、濕度等相關(guān)；③與栽培方式如覆土厚度相關(guān)；④不見原基出菇的成菇率高，但其發(fā)生率更低；⑤同一菌株同一田塊兩種出菇方式都會(huì)發(fā)生；⑥不見原基出菇往往伴隨假根的形成（圖5）。

圖5 不見原基出菇形成的假根

圖6 營(yíng)養(yǎng)袋內(nèi)形成的原基

兩種出菇方式的機(jī)理尚不明確，對(duì)其研究有利于提高成菇率，可控制出菇的密度和整齊度。實(shí)現(xiàn)對(duì)不見原基出菇方式的控制更有利于實(shí)現(xiàn)工廠化栽培。不見原基出菇形成的假根應(yīng)該是環(huán)境因素引起而不是物種的特點(diǎn)，不應(yīng)作為分類依據(jù)。沙質(zhì)土壤滴灌或噴灌易導(dǎo)致土壤微移動(dòng)，這種土壤移動(dòng)是“見原基出菇”方式中原基因移動(dòng)而懸空死亡的重要原因，提高成菇率可在催菇后進(jìn)行噴霧保濕。

工廠化栽培的難點(diǎn)是通過富營(yíng)養(yǎng)栽培提高單位面積的產(chǎn)量。但富營(yíng)養(yǎng)情況下較難出菇，在栽培中曾發(fā)現(xiàn)營(yíng)養(yǎng)袋內(nèi)形成大量原基（圖6），提示條件適宜的情況下富營(yíng)養(yǎng)栽培是可行的。

致謝 本文圖5、圖6由四川省平昌縣金源生物科技有限公司孫勝先生提供，特此感謝。

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