直接種間電子傳遞對緩解厭氧消化抑制效應的研究進展

司哺春，劉凱強，林新宇，劉志丹，楊改秀，張源輝,3

直接種間電子傳遞對緩解厭氧消化抑制效應的研究進展

司哺春1，劉凱強1，林新宇1，劉志丹1，楊改秀2，張源輝1,3

（1. 中國農(nóng)業(yè)大學水利與土木工程學院農(nóng)業(yè)農(nóng)村部設(shè)施農(nóng)業(yè)工程重點實驗室，環(huán)境增值能源實驗室，北京 100083；2. 中國科學院廣州能源研究所，廣州 510640；3. 美國伊利諾伊大學香檳校區(qū)農(nóng)業(yè)與生物工程系伊利諾伊 IL61801）

厭氧消化是將生物質(zhì)廢棄物進行資源化利用的有效途徑之一。然而，復雜的原料性質(zhì)以及反應器高負荷的運行條件會使厭氧消化過程產(chǎn)生多種抑制效應，易導致反應器運行不穩(wěn)定，產(chǎn)氣效率低等問題。因此，提升厭氧消化反應器運行性能、減緩抑制效應成為當前的研究熱點。區(qū)別于以氫氣和甲酸為媒介的間接種間電子傳遞（Mediated Interspecies Electron Transfer，MIET）過程，微生物間的直接種間電子傳遞（Direct Interspecies Electron Transfer，DIET）能夠在菌群間直接進行電子轉(zhuǎn)移，傳遞效率更高。DIET的建立有助于強化厭氧反應的穩(wěn)定性，提高反應效率，減緩抑制效應?；诖耍撐目偨Y(jié)了DIET的研究進展，分析了主要的種間電子傳遞機制，探討了DIET對不同類型抑制效應的緩解作用，歸納了DIET潛在微生物的富集效果；在此基礎(chǔ)上展望了DIET在減緩厭氧消化抑制效應方面的重點研究方向和應用前景。

發(fā)酵；反硝化；抑制效應；微生物群落；導電材料；直接種間電子傳遞

0 引 言

據(jù)統(tǒng)計，世界范圍內(nèi)每年大約產(chǎn)生1 460億t的生物質(zhì)廢物[1]。中國每年產(chǎn)生約21億t 農(nóng)林業(yè)廢棄物和38億t畜禽糞污[2]，但綜合利用率不到60%[3]。廢棄物的不當處理會造成嚴重的環(huán)境污染，包括地表和地下水污染、氣味問題和病原體的傳播[4]。

厭氧消化是一種經(jīng)濟、有效的生物質(zhì)廢物處理和能源生產(chǎn)技術(shù)，由不同微生物共同參與完成產(chǎn)甲烷過程，其反應效率依賴于系統(tǒng)中不同微生物間的協(xié)同作用[5]。高濃度廢水、城市生活垃圾、剩余污泥、畜禽糞污、農(nóng)業(yè)廢棄物、微藻和餐廚垃圾等都可以通過厭氧消化進行能源轉(zhuǎn)化[6-7]。但復雜的原料性質(zhì)以及反應器運行條件的變化會導致厭氧消化過程的多種抑制效應。例如廢水中的氮雜環(huán)化合物、苯酚、呋喃等[8]，高有機負荷（Organic Loading Rate，OLR）運行條件下的中間產(chǎn)物揮發(fā)性脂肪酸（Volatile Fatty Acid，VFA）的積累[9]，含氮物質(zhì)轉(zhuǎn)化導致的氨氮積累[10]等都會對厭氧微生物產(chǎn)生強烈的抑制作用，從而導致厭氧消化無法順利進行[11]。

厭氧消化過程中，互養(yǎng)微生物產(chǎn)乙酸細菌和產(chǎn)甲烷古菌之間的電子傳遞過程由于存在熱力學壁壘的問題，被認為是厭氧消化過程的限速步驟[12]。近年來的研究表明，在厭氧消化過程中，DIET（Direct Interspecies Electron Transfer）是一種比H2或甲酸介導的MIET更有效的種間電子傳遞機制[13]，可加強互養(yǎng)微生物間的“交流”，促進反應自發(fā)進行，從而達到促進厭氧消化的目的[14]。因此，在厭氧消化過程中通過加入導電材料或乙醇，可促進DIET機制建立，進而緩解厭氧消化的抑制效應，強化微生物菌群間的互養(yǎng)代謝過程[15-16]，保證了厭氧消化過程的穩(wěn)定進行[17]。

目前已有文獻探究通過建立DIET來提升厭氧消化性能，但與基于DIET減緩厭氧消化抑制效應的相關(guān)綜述還未有報道。因此，本文旨在總結(jié)種間電子傳遞機制，回顧DIET的研究進展；探討DIET對抑制效應的緩解作用，總結(jié)分析DIET潛在功能微生物群落的富集效果；最后基于以上內(nèi)容提出DIET緩解厭氧消化抑制效應的作用機理探究和工程化應用所面臨的挑戰(zhàn)。

1 種間電子傳遞機制

傳統(tǒng)厭氧發(fā)酵理論認為復雜有機物在被轉(zhuǎn)換為單碳化合物（CO2、甲酸、甲醇、甲胺）和乙酸之前不能產(chǎn)生甲烷，厭氧消化的關(guān)鍵步驟是電子產(chǎn)生與消耗過程[18]。如圖1a所示，MIET是指依賴于H2或甲酸作為載體在互養(yǎng)菌群之間進行的電子傳遞。氫化酶或甲酸脫氫酶的存在對互養(yǎng)微生物通過胞外電子傳遞進行鏈式生化反應是必不可少的。VFAs和乙醇氧化產(chǎn)生的質(zhì)子被氫化酶還原成氫氣，或被甲酸脫氫酶還原成二氧化碳，然后，氫和甲酸將電子從供體（乙酸菌）轉(zhuǎn)移到受體（甲烷菌）。H+/H2的氧化還原電位較低，當與其他氧化還原介質(zhì)發(fā)生耦合反應時，其在標準條件下反應的自由能通常為正值[19]。以丙酸產(chǎn)甲烷過程為例，產(chǎn)乙酸細菌將丙酸氧化成乙酸和氫氣（CH3CH2COO-+ 3H2O → CH3COO-+HCO3-+ H++ 3H2），在標準生化條件下，這個反應過程存在能量壁壘（Δ=76.0 kJ/mol）[20]，進而限制了電子傳遞效率[21]。傳統(tǒng)的MIET機制對電子載體濃度極為敏感，例如，只有當氫營養(yǎng)型產(chǎn)甲烷菌通過消耗H2，使H2壓力維持在較低水平（<1.013 25 Pa）時，MIET過程才能持續(xù)進行[22]。甲酸亦可替代H2作為電子載體，參與電子傳遞過程[23-25]。盡管氫氣的擴散系數(shù)更高，但甲酸介導的MIET具有更高的傳質(zhì)速率[26]。以菲克定律為基礎(chǔ)的擴散模型表明，種間甲酸傳遞比種間氫氣傳遞（Interspecific H2Transfer，IHT）的產(chǎn)甲烷速率高100倍[21]。然而，許多產(chǎn)甲烷古菌在純培養(yǎng)基中不能利用甲酸，種間氫氣傳遞被認為是主要的電子傳遞機制[26]。

2005年，麻省大學Lovley團隊研究發(fā)現(xiàn)，富集在Fe(III)氧化物表面的的菌毛具有良好的導電性，可以將電子直接轉(zhuǎn)移到Fe(III)氧化物中[27]。2010年，該團隊首次證明了電活性微生物和之間存在DIET[28]，并在處理啤酒廢水的上流式厭氧污泥反應器中驗證了顆粒污泥中DIET的形成[29]。Rotaru等[18,30]進一步證實了和常見的產(chǎn)甲烷菌（如、等）能夠進行DIET。如圖 1b所示，互養(yǎng)微生物之間可通過導電菌毛（electrically conductive pili，e-pili）和細胞色素蛋白(c-type Cytochromes，OmcS)進行DIET[16]。通過e-pili和OmcS，細菌可以直接將中間產(chǎn)物產(chǎn)生的電子轉(zhuǎn)移到產(chǎn)甲烷古菌，還原CO2產(chǎn)生甲烷?；ヰB(yǎng)微生物之間也可通過導電材料建立DIET，且機制與導電材料尺寸和結(jié)構(gòu)相關(guān)。如果導電材料的尺寸小于微生物細胞，那么導電材料只能附著在單個細胞上（見圖1e）；而大面積導電碳材料可以在其表面附著多種微生物菌群，進而提供流暢的電子通道（見圖1c）。缺乏e-pili或OmcS，微生物之間無法通過生物連接建立DIET，但添加導電材料后，如圖1d，1f所示，導電材料可以充當電子通道，促進微生物間建立DIET。炭基導電材料（生物炭[2,31]、顆?；钚蕴縖32-33]、石墨烯[34]、碳布[35]等）和鐵基導電材料（磁鐵礦[36-37]、赤鐵礦[38]、針鐵礦[39]以及零價鐵[40]等）具有高導電性、化學穩(wěn)定性、輕質(zhì)和成本較低等優(yōu)勢，被廣泛用于促進厭氧消化過程中DIET的建立。

DIET可通過e-pili和OmcS或結(jié)合生物和非生物電子傳遞元件，如導電材料等進行電子傳遞。與MIET相比，DIET具有以下優(yōu)勢：1）通過反應-擴散-電化學模型計算得到DIET（44.9×103cp-1s-1）的電子傳遞速率是IHT（5.24×103 cp-1s-1）的8～9倍[26]，可加速菌群間的互養(yǎng)代謝過程，提高厭氧消化產(chǎn)氣效率；2）DIET不依賴于介質(zhì)轉(zhuǎn)移電子，無需酶的催化，避免了不必要的能量損耗；3）在DIET過程中，電子直接轉(zhuǎn)移到產(chǎn)甲烷菌，避免了擴散到周圍環(huán)境的損失。

圖1 種間電子傳遞機制

2 DIET對厭氧消化抑制效應的緩解作用

在厭氧消化過程中，微生物群落通常會受到氨氮積累、高濃度硫化物等無機離子以及VFAs積累、芳香族類等難降解有機物引起的抑制作用。目前研究表明微生物菌群間DIET的建立對厭氧消化的抑制效應可起到顯著的緩解作用（表1、表2）。其緩解抑制效應的主要結(jié)果和相應的機制將在下文進行具體闡述。

2.1 無機離子抑制作用的緩解

在厭氧消化過程中，有機態(tài)氮（蛋白質(zhì)、尿素、核酸等）轉(zhuǎn)化成無機態(tài)氨氮。但是由于厭氧微生物生長緩慢，只有少量氨氮可被細胞利用，導致氨氮積累，從而對厭氧消化過程產(chǎn)生抑制作用[41-42]。當氨氮濃度較低時，其作為微生物生長必須的營養(yǎng)物質(zhì)，可以促進微生物生長，但隨著氨氮逐漸累積，當氨氮濃度達到1 500～1 700mg/L[43]時，會對厭氧消化產(chǎn)生較強的抑制性[44-45]。由于產(chǎn)甲烷菌比水解酸化菌的氨氮敏感性更強，因此，高濃度氨氮對產(chǎn)甲烷古菌會產(chǎn)生更顯著的影響[46]。此外，高濃度氨氮抑制產(chǎn)乙酸代謝過程，導致VFAs累積，VFAs濃度超過閾值，使產(chǎn)甲烷菌活性進一步降低，最終形成“抑制的穩(wěn)定狀態(tài)”[44-45,47-48]。近年來，國內(nèi)外初步研究了關(guān)于DIET與厭氧消化氨氮抑制減緩之間的關(guān)系，Zhuang等[43]探究了在不同氨氮濃度下磁鐵礦對厭氧消化的影響，發(fā)現(xiàn)在高氨氮濃度條件下（5 g/L），添加了磁鐵礦的反應器中甲烷效率相比對照組提升58%，磁鐵礦的添加可能促進了乙酸氧化細菌與氫營養(yǎng)型產(chǎn)甲烷古菌之間建立了DIET。類似的，Lee等[49]也發(fā)現(xiàn)在氨氮抑制情況下（6.5g/L），添加磁鐵礦可顯著縮短厭氧消化的延滯期（21%），強化有機酸的轉(zhuǎn)化。Lu等[50]則認為在氨氮濃度逐漸升高的情況下，添加磁鐵礦可促進氫營養(yǎng)型產(chǎn)甲烷菌和產(chǎn)酸菌之間的DIET，進而提升厭氧消化效果。在氨氮逐漸累積的高濃度厭氧消化環(huán)境中，DIET的建立為緩解氨氮抑制，提升反應效率提供了可能。

硫酸鹽是含金屬廢水、紡織廢水、制藥廢水等廢水中的典型成分。硫酸鹽將促進硫酸鹽還原菌的富集，進而硫酸鹽還原菌將會與產(chǎn)甲烷菌競爭電子，同時硫酸鹽的還原產(chǎn)物硫化氫會抑制微生物菌群間的互養(yǎng)代謝，從而影響厭氧消化的性能。DIET機制的建立可以緩解厭氧消化過程中硫酸鹽、硫化氫等對產(chǎn)甲烷過程產(chǎn)生的抑制效應，強化甲烷生成過程。Li等[51]在含硫酸鹽廢水的厭氧消化過程中添加不銹鋼，發(fā)現(xiàn)DIET對電子供體的動力學優(yōu)勢比IHT高108倍，添加不銹鋼條件下反應器的平均產(chǎn)氣量是對照組的4.5倍。Jin等[37]在含硫酸鹽廢水的厭氧消化過程中添加磁鐵礦，發(fā)現(xiàn)磁鐵礦組c型細胞色素的濃度為113.54 nmol/L，約為對照組的一半，表明通過磁鐵礦建立的DIET取代了c型細胞色素，而且，F(xiàn)e(III)還原菌在磁鐵礦表面富集，并與產(chǎn)甲烷菌和建立了DIET通道，使含硫廢水甲烷產(chǎn)量提高了3～10倍，進而推斷DIET是強化甲烷生成和硫酸鹽去除的主要機制。

2.2 有機物抑制作用的緩解

當OLR過高時，VFAs積累、pH下降，厭氧消化產(chǎn)甲烷過程被抑制[35]。在高OLR的反應器中加入碳布[52]、GAC[53]、磁鐵礦[54]、生物炭[55-56]等導電材料，可促進產(chǎn)電細菌和產(chǎn)甲烷古菌之間建立DIET，形成高效的電子傳遞通道，增強微生物間的協(xié)同代謝。DIET功能微生物代謝活性的提高可緩解VFAs積累，提升產(chǎn)甲烷效率，維持厭氧反應器在高OLR下的穩(wěn)定運行（見表2）。例如，丙酸在厭氧消化過程中轉(zhuǎn)化效率低，容易積累，易導致抑制效應。Cruz等[57]探究了添加微米級磁鐵礦對丙酸鹽厭氧消化的影響，通過使用共聚焦激光掃描顯微鏡(Confocal Laser Scanning Microscopy，CLSM)與熒光原位雜交（Fluorescence In Situ Hybridization，F(xiàn)ISH）結(jié)合方法顯示了聚集微生物的存在，古菌總是靠近細菌。進一步推測添加磁鐵礦促進了DIET機制的建立，進而提升了丙酸的降解效率。乙酸是厭氧消化過程中重要的中間代謝產(chǎn)物，Baek等[5]在進行乳制品厭氧消化處理時，添加磁鐵礦可通過建立DIET促進二氧化碳還原和乙酸分解，提高甲烷產(chǎn)量。推測主要是進行了基于DIET的互養(yǎng)乙酸氧化產(chǎn)甲烷途徑[58]。與乙酸營養(yǎng)型產(chǎn)甲烷菌相比，互養(yǎng)乙酸氧化產(chǎn)甲烷菌更能適應高濃度VFA的脅迫[59]。Zhuang等[43]研究發(fā)現(xiàn)，與對照組相比，添加磁鐵礦納米顆?？纱龠M互養(yǎng)乙酸氧化菌和產(chǎn)甲烷菌之間建立DIET機制，從而使乙酸產(chǎn)甲烷速率提高36%～58%。

芳香族化合物結(jié)構(gòu)穩(wěn)定，在厭氧環(huán)境中不易分解，且對微生物毒性很強。水解酸化細菌與產(chǎn)甲烷古菌受到毒性抑制，難以進行產(chǎn)甲烷過程。通過建立DIET，促進了互養(yǎng)微生物代謝活性，可進一步強化芳香族化合物的分解利用。Usman等[8]在厭氧消化處理組分復雜的水熱液化廢水過程中發(fā)現(xiàn)，添加GAC可通過富集DIET潛在功能微生物促進含氮有機物和芳香族化合物的降解。苯甲酸脂是芳香族化合物在厭氧消化過程常見的中間體，Zhuang等[60]發(fā)現(xiàn)通過添加赤鐵礦和磁鐵礦可使苯甲酸酯的降解率分別提高25%和53%。Zhuang等[61]發(fā)現(xiàn)，在厭氧消化反應器中添加赤鐵礦和磁鐵礦可能是由于促進互養(yǎng)苯甲酸酯氧化菌和硫酸鹽還原菌之間建立了DIET，進而使苯甲酸鹽的降解率分別提高了81.8%和91.5%。芘是城市污泥中一種常見的多環(huán)芳烴，Li等[62]在城市污泥的厭氧消化過程中添加FeS和磁性碳，通過建立DIET可使芘去除率由40.8%分別提高到77.5%和72.1。

表1 DIET緩解無機離子抑制作用

注：AnDMBRs：厭氧動態(tài)膜生物反應器，AMPTSII：全自動甲烷潛力測試系統(tǒng)，CSTR：全混流反應器，UASB：上流式厭氧污泥床，ASBR：厭氧序批反應器。

Note: AnDMBRs: anaerobic dynamic membrane bioreactor, AMPTSII: automated methane potential testing system, CSTR:continuous stirred tank reactor, UASB: upflow anaerobic sludge blanket, ASBR: anaerobic sequencing batch Reactor.

2.3 DIET潛在功能微生物的富集

雖然大量研究表明DIET可緩解厭氧消化過程中高濃度無機物和有機物的抑制作用，但如何針對性的建立微生物間的DIET，確認DIET的存在，從而揭示緩解厭氧消化抑制效應的機制，需要對DIET功能和潛在功能微生物進一步深入研究。到目前為止，有直接且充分證據(jù)證明具有DIET功能的細菌包括、以及，具有DIET功能的古菌為、以及和[28,68-71]。作為MIET研究最深入的微生物模型之一，2020年，Walker等[71]的研究表明其可通過導電菌毛建立DIET。因此，DIET機制可能是進行MIET微生物群落的一種選擇，這表明能夠參與DIET的微生物群落可能比已確認的更廣泛。

在厭氧消化過程中添加導電材料，會富集許多具有DIET潛力的電活性微生物。圖2所示微生物菌群中，和是水解過程中主要的菌群；、、和是酸化過程中主要的菌群[72]。除中的產(chǎn)電細菌可與產(chǎn)甲烷古菌和建立DIET機制外，以上幾類菌門中的許多產(chǎn)電細菌也具有與乙酸營養(yǎng)型或氫營養(yǎng)型產(chǎn)甲烷菌建立DIET機制的潛力。

供電子細菌的富集，可被認為是微生物菌群間建立DIET的有效證據(jù)。Lin等[34]研究發(fā)現(xiàn)，以乙醇為發(fā)酵底物，在厭氧消化過程中添加石墨烯后，產(chǎn)電細菌和的豐度提高，氫營養(yǎng)型產(chǎn)甲烷菌和占據(jù)了主導地位，進而推測電子供體細菌、以及電子受體古菌、之間建立了DIET，增強了厭氧消化的性能。Guo等[23]通過在厭氧反應器中添加GAC，發(fā)現(xiàn)、、和28以及、和在GAC生物膜上得到富集，這些與DIET相關(guān)的電活性微生物擴展了DIET潛力微生物范圍。Lee等[73]在以乙酸為底物的厭氧消化反應器中添加GAC，觀察到除得到富集外，、以及氫營養(yǎng)型產(chǎn)甲烷菌和的豐度也得到提高，進一步推測除外，其他細菌也具有參與DIET的潛力。

圖2 DIET潛在功能微生物群落

厭氧消化過程在受到較強的抑制作用下，不能有效地降解復雜有機物[74]。當豐度較低時，DIET被推測可能是其他重要的細菌參與完成的。Capson等[10]發(fā)現(xiàn)，高氨氮濃度的食品廢棄物在進行厭氧消化時，未檢測到電活性細菌，所以推測厭氧消化抑制情況下參與DIET的細菌可能是和。Lei等[35]在處理焚燒垃圾滲濾液的UASB反應器中添加碳布，推測和能夠以碳布為電子通道，與產(chǎn)電細菌、和進行DIET。Zhang等[13]在厭氧消化處理餐廚垃圾油菜籽的試驗中發(fā)現(xiàn)，GAC表面富集了大量，進一步說明具有參與DIET的潛力。

胞外電子轉(zhuǎn)移到Fe(III)氧化物的機制通常與電子通過DIET轉(zhuǎn)移到產(chǎn)甲烷菌的機制密切相關(guān)。與一樣，也是一種Fe(III)還原菌，可在添加木屑生物炭的條件下得到富集，因此推測參與了DIET[9]。Wang等[55]在厭氧消化過程中添加木屑生物炭，發(fā)現(xiàn)的豐富度明顯增加，屬于Fe(III)還原菌屬，具有生成菌毛的pilA基因，而且與已確定能進行DIET的同時得到了富集。也屬于Fe(III)還原菌，在磁鐵礦存在的條件下在厭氧消化反應器中被富集，因此，推測通過建立DIET進行共養(yǎng)代謝[37]。Zhao等[16]將剩余污泥與乙醇發(fā)酵液混合進行厭氧消化，發(fā)現(xiàn)污泥電導率提高約3-5倍，且Fe(III)還原菌和具有高豐度導電菌毛的被富集，因此推斷和之間建立了DIET。Jia等[64]在處理啤酒廢水的厭氧消化反應器中添加碳布發(fā)現(xiàn)細菌得到富集。表明也具有參與DIET的潛力。

3 結(jié)論和展望

厭氧消化抑制效應的緩解，除進行原料預處理、厭氧反應器結(jié)構(gòu)的設(shè)計以及反應參數(shù)的優(yōu)化外，促進微生物菌群間建立DIET是一種具有潛力和前景的方法。

雖然目前已有諸多研究證明了DIET對厭氧消化抑制效應的緩解效果，然而還存在諸多挑戰(zhàn)：1）已經(jīng)證實能進行DIET的互養(yǎng)微生物種類單一。不同于純培養(yǎng)體系，目前大部分研究是在復雜生物質(zhì)廢物厭氧消化過程中基于導電材料對微生物的特異性富集來推測能進行胞外電子傳遞的微生物是否參與了DIET，其結(jié)果還需要進一步驗證。2）已有研究推測氫營養(yǎng)型產(chǎn)甲烷菌也參與了DIET，然而在氫營養(yǎng)型產(chǎn)甲烷菌主導的反應系統(tǒng)中，IHT也會被促進，即使存在DIET，也難以驗證DIET在厭氧消化過程的主導地位。3）在厭氧消化過程中加入導電材料或乙醇等可以促進微生物間建立DIET，但相關(guān)機理仍需進一步探究，特別是如何準確論證DIET的形成以及DIET對抑制效應的緩解作用。4）考慮到工程應用時，導電材料如何在大型反應器中持續(xù)穩(wěn)定的發(fā)揮作用以及導電材料的成本和回收等仍是需要解決的難題。

因此，進一步的研究可以考慮但不局限于以下幾個方面：

1）通過長期連續(xù)試驗驗證，基于16S rRNA 基因序列分析，結(jié)合蛋白質(zhì)組學和宏基因技術(shù)，確定厭氧體系中參與抑制物降解以及能夠進行DIET的微生物，拓展DIET功能微生物的數(shù)據(jù)庫。此外，基于復雜模型的定量化對比，例如厭氧消化1號模型（Anaerobic Digestion Model No. 1，ADM1），也將有助于揭示DIET緩解厭氧消化抑制機制。

2）厭氧消化反應器中添加導電材料，在探究DIET減緩厭氧消化抑制作用的同時，也要全面考慮導電材料比表面積、表面粗糙度和吸附性能等特性對厭氧消化的促進作用，建立導電材料物化特性和厭氧強化機理以及DIET機制的對應關(guān)系。

3）目前實驗室規(guī)模研究已經(jīng)證明多種導電材料能夠促進微生物間的DIET形成，減緩抑制效應。然而，在實際工程中，導電材料的經(jīng)濟技術(shù)可行性、環(huán)保性的評價將成為重要的考量因素。因此，進一步對厭氧反應器內(nèi)導電材料添加的經(jīng)濟技術(shù)分析以及生命周期評估的研究將極為重要。

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Research progress of the relief of anaerobic digestion inhibitions based on direct interspecies electron transfer

Si Buchun1, Liu Kaiqiang1, Lin Xinyu1, Liu Zhidan1, Yang Gaixiu2, Zhang Yuanhui1,3

(1.(2),,,,,100083,; 2.,.510640,; 3.,,IL 61801,)

Anaerobic digestion is one of the most effective technologies to valorize biomass waste for bioenergy production. However, complex characteristics of raw materials and varied operating conditions of the reactor resulted in multiple inhibitions during anaerobic digestion. Therefore, improving the performance of anaerobic reactors and reducing the inhibitory effects have attracted increasing attentions. Unlike mediated interspecies electron transfer (MIET) by hydrogen or formic acid, direct interspecies electron transfer (DIET) can directly transfer electrons among microbes. Establishing DIET could enhance the stability of digestion, improve reaction efficiencies, and relieve inhibitory effects during anaerobic digestion. However, DIET is still in its infancy stage. To this end, this study aim to summarize the main mechanisms of interspecific electron transfer, in particular, review and discuss the effects of DIET on relief of inhibitions of anaerobic digestion. DIET can be established via e-pili, OmcS and conductive materials between syntrophic bacteria and methanogens. Compared with electron transfer via MIET, DIET shows several advantages. Electron transfer rate of DIET is 8-9 times that of MIET. In addition, DIET does not depend on the medium to transfer electrons, which indicates that it has no requirement of hydrogenase or formate dehydrogenases, thus avoiding unnecessary energy loss. What’s more, during the DIET, electrons are directly transferred between bacteria and methanogens, and it avoids the loss of electrons caused by the diffusion to surrounding environment. Further, the effects of DIET on different inhibitions were summarized and discussed. Present studies indicated the inhibitions caused by nitrogen heterocyclic compounds, aromatic organics and furans, and accumulated volatile fatty acids could be significantly relived with the establishment of DIET. In addition, DIET was also proved could effectively improve the anaerobic digestion with inorganic inhibitors, high content of ammonia and sulfur-containing compounds. Adding conductive materials or ethanol could stimulate the establishment of DIET and enrich DIET related microbes, so as to alleviate the inhibitory effects and strengthen the activities of microbial communities. The enriched DIET related microorganisms were also summarized in this review. So far, microbes includedandwere confirmed involved in DIET. Other syntrophic bacteria, such as,,,,,,,,,,,,andshowed a potential to establish DIET with Methanogens. Lastly, future researches and application prospects of DIET for relieving inhibitions of anaerobic digestion were proposed. 1) The direct evidence for verifying DIET related microbes should be provided. Most of present studies proposed DIET related microorganisms were based on indirect evidence, such as the specific enrichment by conductive materials and improved methane production; 2) It has been speculated that hydrogenotrophic methanogens also participated in DIET. However, MIET will also be promoted in the system dominated by hydrogenotrophic methanogens. The roles of DIET and MIET during anaerobic digestion with inhibitors are desperate to reveal; 3) In depth mechanisms of relieved inhibitions via addition of conductive materials need further exploration. There is an urgent requirement for building up the corresponding relationship between physicochemical properties of additives and establishment of DIET; 4) The techno-economic analyses and environmental evaluations of conductive materials should be conducted to provide insights for commercial application of DIET enhanced anaerobic digestion.

fermentation; inhibitors; microbial communities; conductive materials; direct interspecies electron transfer

司哺春，劉凱強，林新宇，等. 直接種間電子傳遞對緩解厭氧消化抑制效應的研究進展[J]. 農(nóng)業(yè)工程學報，2020，36(24)：227-235.doi：10.11975/j.issn.1002-6819.2020.24.027 http://www.tcsae.org

Si Buchun, Liu Kaiqiang, Lin Xinyu, et al. Research progress of the relief of anaerobic digestion inhibitions based on direct interspecies electron transfer[J]. Transactions of the Chinese Society of Agricultural Engineering (Transactions of the CSAE), 2020, 36(24): 227-235. (in Chinese with English abstract) doi：10.11975/j.issn.1002-6819.2020.24.027 http://www.tcsae.org

2020-09-07

2020-11-27

國家自然科學基金青年項目（51806243）

司哺春，博士，副教授，主要從事厭氧發(fā)酵產(chǎn)甲烷抑制解除及過程強化研究，暗發(fā)酵用于農(nóng)業(yè)廢棄物產(chǎn)氫及酸化產(chǎn)物調(diào)控以及厭氧高效反應器的設(shè)計與優(yōu)化。Email：sibuchun@cau.edu.cn

10.11975/j.issn.1002-6819.2020.24.027

S2

A

1002-6819(2020)-24-0227-09