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原生生物
原生生物

原生生物

物種分類

原生生物（學名：Protist）係一類真核生物，主要包含有細胞核嘅單細胞生物，唔係動物、植物同真菌。雖則歸為一類，並唔係因為佢哋之間有祖先關連，只係方便研究，未清楚關係之前，暫時擺埋一齊。佢哋之間差異極大，細胞結構、繁殖同生活史，可以極之唔同。如今種種生物，視為原生生物演化出來。 原生生物喺歷史上曾經俾人當做一個獨立嘅分類學 界，叫做 原生生物界 或者 原生生物總界。隨住系統發生學分析同電子顯微鏡研究嘅興起，原生生物界作為一個正式分類單元嘅用法就逐漸冇人再用。而家嘅分類入面，原生生物分散喺幾個真核生物演化支，叫做超門，好似古蟲界 （光合自養生物，包括陸地植物）、SAR、後鞭毛生物（包括真菌同動物）、變形蟲同「Excavata」。

原生生物代表咗極其龐大嘅遺傳同生態多樣性，喺所有環境入面，包括極端棲息地。佢哋嘅多樣性，大過所有其他真核生物，只係喺近幾十年透過研究環境DNA先至發現，而且重喺完全描述嘅過程入面。佢哋存在於所有生態系統，作為生物地球化學循環同食物網嘅重要組成部分。佢哋豐富而且普遍噉存在於各種主要係單細胞嘅形態，呢啲形態獨立噉進化咗好多次，好似自由生活嘅藻類、變形蟲同黏菌，又或者作為重要嘅寄生生物。加埋一齊，佢哋構成嘅生物量係動物嘅兩倍。佢哋展現出唔同類型嘅營養方式（好似光合營養、吞噬營養或者滲透營養），有時會結合埋一齊（喺混合營養入面）。佢哋呈現出多細胞動物、真菌或者陸地植物冇嘅獨特適應性。對原生生物嘅研究叫做原生生物學。

形態學 (A) 同遺傳學 (B) 角度睇真核生物總體多樣性嘅差異。原生生物喺 DNA條碼 分析中佔據主導地位，但係喺編目物種中僅佔少數。[31]

原生生物嘅物種多樣性俾傳統方法嚴重低估，傳統方法基於形態特徵嚟區分物種。已描述嘅原生生物物種數量非常低（範圍由 26,000[32] 到超過 76,000）[c]，相比之下，植物、動物同真菌嘅生物多樣性歷史悠久，生物學上都好出名而且研究充分。預測嘅物種數量差異都好大，範圍由 1.4×105 到 1.6×106，喺幾個群體中，預測嘅物種數量任意噉增加一倍。呢啲預測大多數都係高度主觀嘅。分子技術，例如 環境 DNA條碼，揭示咗大量未描述嘅原生生物，佢哋佔咗大多數真核生物序列或者操作分類單元 (OTU)，令植物、動物同真菌相形見絀。[31] 因此，原生生物俾人認為喺真核生物多樣性中佔據主導地位。[34]

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原生生物嘅進化關係已經通過分子系統發生學、測序完整嘅基因組同轉錄組，以及電子顯微鏡對鞭毛裝置同細胞骨架嘅研究嚟解釋。原生生物嘅新主要譜系同新嘅生物多樣性繼續被發現，導致真核生物生命樹發生巨大變化。最新嘅真核生物分類系統唔承認正式嘅分類階元（界、門、綱、目……），而係只承認相關生物嘅演化支，令分類喺長期內更加穩定，並且更容易更新。喺呢個新嘅支序分類學方案中，原生生物被劃分為各種分支，非正式噉命名為超門。大多數光合真核生物都屬於 Diaphoretickes 演化支，其中包括超門 古蟲界（包括植物）同 TSAR（包括 Telonemia、不等鞭毛類、囊泡藻 同 有孔蟲），以及門 Cryptista 同 Haptista。[14] 動物同真菌屬於 Amorphea 超門。


Diaphoretickes 包括差唔多所有光合作用真核生物。喺呢個演化支入面，TSAR 超門聚集咗大量嘅原生生物多樣性。TSAR 最基部分支成員係 Telonemia，一個細小（七個物種）嘅門，佢哋係唔出名嘅吞噬性捕食性鞭毛蟲，喺海洋同淡水環境度搵到（但係佢亦都有可能係 Haptista 嘅姊妹演化支，即係唔形成假設嘅 TSAR 演化支[49]）。佢哋同剩低嘅三個演化支共享一啲細胞相似性：有孔蟲、囊泡藻 同 不等鞭毛類，統稱為 SAR。[50] Diaphoretickes 內另一個高度多樣化嘅演化支係 古蟲界，佢包含 陸地植物 同各種藻類。此外，兩個較細嘅群體，Haptista 同 Cryptista，都屬於 Diaphoretickes。[8]

不等鞭毛類，亦都叫做 Heterokonta，其特徵係存在兩條纖毛，其中一條纖毛帶有好多短嘅稻草狀毛 (羽鞭毛)。佢哋包括一個光合自營生物演化支同埋好多個異營生物演化支，幾乎喺所有棲息地都有。不等鞭毛類包括兩個通常得到良好支持嘅演化支，Bigyra 同 Gyrista，雖然 Bigyra 嘅單系群性受到質疑。[51] 喺 Bigyra 同 Gyrista 之外分支嘅係一種神秘嘅異養鞭毛蟲，Platysulcus tardus。[51] 大部分異養不等鞭毛類嘅多樣性仍然未被描述，幾乎完全從遺傳序列譜系（叫做 MASTs (MArine STramenopiles)）度得知，[51] 其中只有少數物種俾人描述過。[52][53]

門 Gyrista 包括光合金藻門或者 Heterokontophyta （超過二萬三千個物種），[42] 佢哋包含起源於紅藻嘅葉綠體。喺呢啲之中有好多藻類譜系，佢哋包含廣泛嘅結構同形態。三個最多樣化嘅金藻綱係：矽藻，被包裹喺二氧化矽細胞壁（殼）入面嘅單細胞或者群體生物，佢哋展現出廣泛唔同嘅形狀同裝飾，負責全球產生嘅大部分氧氣，並且構成海洋浮游植物嘅大份；[14][54] 褐藻，絲狀或者「真正」多細胞（具有分化組織）巨型藻類，構成咗好多溫帶同寒冷海洋生態系統嘅基礎，例如海帶森林；[55] 同 金黃藻，單細胞或者群體鞭毛蟲，主要存在喺淡水棲息地。[56] 喺 Gyrista 入面，金藻門嘅姊妹演化支係主要滲透營養同絲狀嘅 假菌（超過一千二百個物種），[57] 佢哋包括三個唔同嘅譜系：寄生性嘅 卵菌 或者水黴菌（例如，致病疫黴菌，愛爾蘭大饑荒背後嘅病原體），佢哋包含咗大多數假菌物種；多樣性冇咁高嘅非寄生性 絲狀壺菌，佢哋維持住類似真菌嘅生活方式；同 Bigyromonadea，一個食菌性或者食真核生物吞噬生物群體。[51] 一個細群體嘅類似太陽蟲嘅異養變形蟲，太陽藻綱，位置唔確定，可能喺金藻門入面，亦都有可能係佢嘅姊妹分類單元。[58]

門 Bigyra 係一個完全異養生物嘅集合，其中大多數係自由生活嘅。佢包括 迷宮藻綱，喺佢哋之中有單細胞變形蟲吞噬生物、混合營養生物，同埋類似真菌嘅絲狀異養生物，佢哋創造黏液網絡嚟移動同吸收營養，以及一啲寄生蟲同少量有殼變形蟲 (Amphitremida)。Bigyra 入面仲包括 領鞭毛蟲，食菌性鞭毛蟲，佢哋食細菌，以及密切相關嘅 Placidozoa，Placidozoa 由幾個異養鞭毛蟲群體組成（例如，深海嗜鹽 Placididea），以及叫做 Opalinata 嘅腸道共生生物（例如，人類寄生蟲 Blastocystis，同非常唔尋常嘅 多核纖毛蟲，由具有大量細胞核同纖毛嘅巨型細胞組成，最初俾人錯誤分類做纖毛蟲）。[51]


囊泡藻 (Alveolata) 嘅特徵係存在皮質泡囊，細胞質囊泡位於細胞膜之下，生理功能未知。[35]:{{{1}}} 喺佢哋之中有三個最出名嘅原生生物群體：頂複門、甲藻同纖毛蟲。纖毛蟲綱 (Ciliophora) 係一個高度多樣化（超過八千個物種），可能係研究得最透徹[14] 嘅原生生物群體。佢哋大多數係自由生活嘅微生物，特徵係大型細胞，表面覆蓋住成排纖毛，並且包含兩種細胞核，小核同大核（例如，草履蟲，一個模式生物）。[59] 自由生活嘅纖毛蟲通常係微生物食物網中嘅頂級異養生物同捕食者，以細菌同較細嘅真核生物做嘢食，存在喺各種生態系統入面，雖然少數物種係盜食質體。其他嘅係好多動物嘅寄生蟲。[60] 纖毛蟲喺囊泡藻嘅進化中處於基部位置，連同少數幾種具有兩條纖毛嘅異養鞭毛蟲，統稱為 Colponemida。[61]

剩低嘅囊泡藻俾人歸類喺 Myzozoa 演化支之下，佢哋嘅共同祖先通過次級內共生作用由紅藻度獲得葉綠體。[62] Myzozoa 嘅一個分支包含頂複門同佢哋最接近嘅親戚，一個叫做 Chrompodellida 嘅細鞭毛蟲演化支，喺呢個演化支入面，光合自養同異養鞭毛蟲（分別叫做 Chromerida 同 Colpodellida）喺進化上互相混合。[62] 相反，頂複門 (Apicomplexa) 係一個龐大（超過六千個物種）而且高度專業化嘅專性寄生蟲群體，佢哋全部都二次性噉冇咗光合作用能力（例如，惡性瘧原蟲，瘧疾嘅成因）。佢哋嘅成蟲階段通過細胞膜由寄主度吸收養分，而且佢哋喺寄主之間通過孢子蟲繁殖，孢子蟲展現出一個由非光合葉綠體進化而嚟嘅細胞器複合體（頂質體）。[63][35]:{{{1}}}

Myzozoa 嘅另一個分支包含甲藻同佢哋最接近嘅親戚，Perkinsidae (Perkinsozoa)，一個細群體（廿六個物種）嘅水生細胞內寄生蟲，佢哋同頂複門類似噉冇咗光合作用能力。[62] 佢哋通過鞭毛孢子繁殖，鞭毛孢子會感染甲藻、軟體動物同魚。[64] 相反，甲藻門 (Dinoflagellata) 係一個高度多樣化（約莫四千五百個物種）[65] 嘅水生藻類群體，佢哋大多數都保留咗佢哋嘅葉綠體，雖然好多譜系都冇咗佢哋自己嘅葉綠體，而係轉為作為異養生物生活，或者由其他來源重新獲得新嘅葉綠體，包括三級內共生同盜食質體。[66] 大多數甲藻都係自由生活嘅，並且構成浮游植物嘅重要組成部分，以及有害藻華嘅主要成因，因為佢哋嘅毒性；一啲以珊瑚嘅共生體形式生活，令珊瑚礁得以形成。甲藻展現出多樣化嘅細胞結構，好似複雜嘅類眼點眼斑、特殊嘅液泡、生物發光細胞器，以及包圍細胞嘅壁，叫做甲殼。[65]

Haptista 同 Cryptista 係兩個相似嘅單細胞原生生物門，之前俾人認為佢哋關係密切，統稱為 Hacrobia。[80] 然而，Hacrobia 嘅單系群性俾人推翻咗，因為呢兩個群體係獨立起源嘅。[81] 分子分析將 Cryptista 放喺古蟲界嘅隔離，形成咗假設嘅「CAM」演化支，而 Haptista 就放喺 Telonemia 同 SAR 演化支隔離（Telonemia 可能係 SAR 嘅姊妹群，形成咗假設嘅 TSAR 演化支，[82] 又或者係 Haptista 嘅姊妹群，形成一個同 SAR 共同嘅姊妹演化支[49][83]）。[83][38]

門 Haptista 包括兩個唔同嘅、帶有礦化鱗片嘅演化支：定鞭藻 同 太陽中心蟲。[14] 定鞭藻 (Haptophyta) 係一個超過五百個現存物種[42] 嘅群體，佢哋係鞭毛或者球狀藻類，已經由次級內共生作用獲得咗葉綠體。佢哋大多數都係海洋生物，構成海洋浮游植物嘅重要組成部分，而且包括 顆石藻，佢哋嘅鈣化鱗片（「顆石」）有助於沉積岩嘅形成以及碳同鈣嘅生物地球化學循環。一啲物種曉得形成有毒藻華。[84] 太陽中心蟲 (Centroplasthelida) 係一個細小（約莫九十五個物種）[85] 但分佈廣泛嘅群體，佢哋係異養太陽蟲型變形蟲，通常都俾帶鱗片嘅黏液覆蓋，構成咗水生棲息地（包括海洋同淡水）底棲食物網嘅重要組成部分。[86]

門 Cryptista 係三個唔同嘅單細胞原生生物群體嘅一個演化支：隱藻、Katablepharidophyta，同埋物種 Palpitomonas bilix。[8] 隱藻（超過一百個物種），亦都叫做隱藻植物，係喺唔同鹽度嘅水生棲息地度搵到嘅鞭毛藻類，佢哋嘅特徵係具有外突細胞器或者擠出器，叫做彈射體。佢哋嘅葉綠體，紅藻起源，包含核藻體，內共生紅藻嘅真核細胞核殘餘物。[87] Katablepharidophyta，隱藻嘅近親，係具有兩條纖毛嘅異養鞭毛蟲，亦都以彈射體做特徵。[80][8] 物種 Palpitomonas bilix 係 Cryptista 入面最基部分支嘅成員，一種海洋異養鞭毛蟲，具有兩條纖毛，但係同剩低嘅成員唔同嘅係，佢冇彈射體。[88]


總體嚟講，原生生物具有典型嘅真核細胞，佢哋遵循生物學中描述嘅嗰啲「更高等」真核生物（動物、真菌同植物）細胞嘅相同原理。[127] 但係，好多原生生物都進化出各種獨特嘅生理適應性，呢啲適應性喺剩低嘅真核生物中係睇唔到嘅，[128] 事實上，原生生物幾乎包含嗮預期喺真核生物中嘅所有廣泛生物學特徵。[34]

原生生物展現出各種各樣嘅食物偏好同埋餵食機制。[8][129] 根據佢哋嘅養分來源，佢哋可以劃分做「自營生物」（生產者）同「異營生物」（消費者）。自營原生生物合成佢哋自己嘅有機化合物，通過光合作用，由無機底物度合成，用光做能量來源；[130]:{{{1}}} 因此，佢哋亦都叫做「光合營養」生物。[131]

異養原生生物獲取由其他生物合成嘅有機分子，而且可以根據佢哋嘅養分大小進一步劃分。嗰啲以可溶分子[130]:{{{1}}} 或者細過 0.5 μm 嘅大分子做嘢食嘅，叫做「滲透營養」生物，[129] 佢哋通過擴散、纖毛孔、細胞膜嘅轉運蛋白同埋一種 內吞作用（即係細胞膜向內凹陷成液泡，叫做 內體）嚟吸收佢哋，內吞作用就叫做 胞飲作用[8] 或者液相內吞作用。[129] 嗰啲以大過 0.5 μm 嘅有機顆粒或者成個細胞做嘢食嘅，叫做「吞噬營養」生物，佢哋通過一種叫做 吞噬作用 嘅內吞作用類型嚟攝取佢哋。[129][130]:{{{1}}} 內吞作用俾人認為係真核生物起源入面最重要嘅進化適應之一，因為佢增加咗潛在嘅食物供應，而吞噬作用就容許咗內共生同線粒體同葉綠體嘅發育。無論係滲透營養生物定吞噬營養生物，內吞作用通常都限於細胞膜嘅特定區域，叫做 胞口，胞口之後可能會係胞咽，胞咽係一個由 微管 支撐嘅特殊管道。[129]

滲透營養原生生物通過膜通道同載體嚟獲取可溶性養分，但係亦都通過唔同類型嘅胞飲作用嚟獲取。巨胞飲作用涉及細胞膜摺疊成褶邊，[132] 產生大型（0.2 到 1.0 μm）液泡。微胞飲作用涉及較細嘅囊泡，通常由 網格蛋白 形成。喺兩種情況下，囊泡都會融合到一個消化液泡或者內體，消化作用喺嗰度發生。[129] 一啲滲透營養生物，叫做「腐生生物」或者「溶養生物」，通過將酶釋放到環境入面並分解有機物，嚟進行體外消化，[8] 將有機物分解做更簡單嘅分子，然後就可以吸收。呢種體外消化具有明顯嘅優勢：佢容許對允許進入細胞嘅物質進行更大嘅控制，從而最大程度噉減少有害物質或者感染嘅攝入。[133]

可能所有真核生物都有滲透營養嘅能力，但係一啲就冇其他獲取養分嘅方法。專性滲透營養生物同腐生生物包括一啲 眼蟲藻、一啲 綠藻、人類寄生蟲 Blastocystis、一啲 Metamonada，[8] 寄生性 鞭毛血原蟲，[134] 同埋似真菌嘅 卵菌 同 絲壺菌。[133]

錐蟲 中胞口-胞咽複合結構嘅結構。食物沿住鞭毛袋嘅口前脊移動，直到到達胞口，並通過胞咽進入細胞，養分估計喺嗰度通過 肌球蛋白 蛋白質運輸，直到佢哋俾囊泡包圍。胞咽由特定組嘅微管支撐。[135]
草履蟲 具有伸縮泡

好多淡水原生生物需要進行滲透調節（即係，去除多餘嘅水份嚟調整離子濃度），因為非鹽水會通過滲透作用由環境大量流入，[142] 同埋通過內吞作用喺食嘢嗰陣時流入。[143] 滲透調節係通過細胞膜嘅主動離子轉運蛋白同埋通過伸縮泡嚟完成嘅，伸縮泡係特殊嘅細胞器，佢哋通過舒張期同收縮期嘅循環，週期性噉排出富含鉀同鈉嘅液體。當細胞放置喺唔同鹽度嘅介質入面嗰陣時，循環就會停止，直到細胞適應。[128]

伸縮泡俾 海綿體 包圍，海綿體係一系列細胞質囊泡或者管道，佢哋由細胞質度慢慢噉收集液體到液泡入面。然後，液泡收縮，並通過一個孔將液體排放到細胞外邊。收縮機制因原生生物而異：喺纖毛蟲入面，海綿體由唔規則嘅小管組成，肌動蛋白絲纏繞喺孔同液泡表面，連同微管；喺大多數鞭毛蟲同變形蟲入面，海綿體由囊泡同小管組成；喺甲藻入面，鞭毛根絲分支形成一個喺液泡周圍嘅收縮鞘（叫做排泄泡）。[143] 位置同數量亦都唔同：單細胞鞭毛藻（隱藻、眼蟲藻、綠藻、金藻、定鞭藻等等）通常喺固定位置具有單個伸縮泡；裸變形蟲有好多細小嘅囊泡，佢哋會融合到一個液泡，然後喺排泄之後再次分裂。海洋或者寄生性原生生物（例如，Metamonada），以及嗰啲具有剛性細胞壁嘅原生生物，就冇呢啲液泡。[142]

自由生活原生生物嘅共識生命週期，包括有性繁殖（紅色箭頭）、無性繁殖（綠色箭頭）、群體階段（藍色），同埋胞囊嘅形成。每個原生生物群體都有唔同嘅有性循環（淺紫色），以及唔同嘅離開群體階段嘅方式。[145]

原生生物展現出好大範圍嘅生命週期同策略，包括唔同形態嘅多個階段，呢啲令佢哋能夠喺大多數環境中蓬勃發展。但係，關於原生生物生命週期嘅大多數知識都係關於模式生物同重要寄生蟲。自由生活嘅未培養原生生物佔大多數，但係關於佢哋生命週期嘅知識仍然支離破碎。[145]

雖然無性繁殖仍然係原生生物中最常見嘅策略，但係有性繁殖亦都係真核生物嘅基本特徵。[149][150] 有性繁殖涉及減數分裂（一種特殊嘅細胞核分裂，容許基因重組）同 配子結合（嚟自兩個親本嘅細胞核嘅融合）。[145] 呢啲過程俾人認為喺 最後共同真核祖先 中已經存在，[151] 佢好可能具有喺兼性（非強制性）基礎上進行有性繁殖嘅能力。[152] 即使係唔再進行有性繁殖嘅原生生物，仍然保留住一組核心嘅減數分裂相關基因，反映出佢哋係由有性祖先傳承落嚟嘅。[153][154] 例如，雖然 變形蟲 傳統上俾人認為係無性生物，但係大多數無性變形蟲可能係最近同獨立噉由有性繁殖變形蟲祖先度出現嘅。[155] 即使喺 20 世紀初，一啲研究人員都將變形蟲入面同染色質絲（游離喺細胞質中嘅 染色質 顆粒）相關嘅現象解釋做有性繁殖。[156]

每個有性循環都涉及配子結合同減數分裂嘅事件，佢哋分別增加或者減少 倍性（即係染色體組嘅數量，用字母 n 表示）。配子結合意味住兩個單倍體 (1n) 繁殖細胞（叫做 配子）嘅融合，產生一個叫做 合子 嘅二倍體 (2n) 細胞。二倍體細胞然後進行減數分裂，產生單倍體細胞。取決於邊啲細胞組成個體或者營養階段（即係通過有絲分裂生長嘅階段），喺自由生活原生生物中觀察到三種可區分嘅有性循環：[145]

喺 單倍體循環 中，個體係單倍體，並且通過有絲分裂分化成單倍體配子。配子融合為一個合子，合子會即刻進行減數分裂，產生新嘅單倍體個體。[145] 一啲 綠藻（即係 團藻目）、好多 甲藻、一啲 Metamonada，同 頂複門 都係噉嘅情況。[143]:{{{1}}}

兩隻纖毛蟲喺接合期間結合埋一齊，通過細胞質橋交換佢哋嘅單倍體細胞核。

喺 雙倍體循環 中，個體係二倍體，並且進行減數分裂，產生單倍體配子，而單倍體配子又會同其他配子融合，形成一個合子，合子發育成一個新個體。[145] 一啲 Metamonada、太陽蟲、好多綠藻、矽藻 同 纖毛蟲，以及動物都係噉嘅情況。[143]:{{{1}}} 纖毛蟲唔係產生配子，而係將佢哋嘅二倍體 小核 分裂成兩個單倍體細胞核，通過接合同另一隻纖毛蟲交換其中一個細胞核，然後將兩個細胞核融合為一個新嘅二倍體細胞核。[60]

喺 單雙倍體循環 中，有兩個世代交替嘅個體。一代係二倍體「無配子體」，佢進行減數分裂，產生單倍體細胞（孢子），孢子發育成另一代，單倍體「配子體」。然後，配子體通過有絲分裂產生配子，配子又融合形成合子，合子發育成無配子體。[145] 好多 有孔蟲 同好多藻類，以及陸地植物都係噉嘅情況。[143]:{{{1}}} 呢個循環有三種模式，取決於一代嘅相對生長同壽命同另一代相比係點：單倍體優勢、二倍體優勢或者世代勢均力敵。褐藻 展現出呢啲模式嘅完整範圍。[157]

自由生活嘅原生生物傾向於喺壓力條件下進行有性繁殖，例如飢餓或者熱衝擊。氧化應激 會導致 DNA損傷，似乎亦都係原生生物誘發性行為嘅重要因素。[146]

病原性原生生物傾向於具有極其複雜嘅生命週期，涉及生物嘅多種形態，其中一啲進行有性繁殖，而其他一啲就無性繁殖。[158] 喺寄主體內食嘢同繁殖嘅階段通常叫做「滋養體」（嚟自Template:Etymology/lang trophos，意思即係：“nutrition”,和 zoia，意思即係：“animals”），但係每個階段嘅名稱因原生生物群體而異。[148] 例如：

喺頂複門入面，單倍體「孢子蟲」釋放到寄主入面，穿透寄主細胞，開始感染並轉化為「裂殖體」，裂殖體生長並無性分裂成大量「裂殖子」（一種叫做「裂殖生殖」嘅裂殖生殖）；每個裂殖子都通過繁殖嚟延續感染。最終，裂殖子分化（「配子生殖」）成雌性（「巨配子母細胞」）同雄性（「微配子母細胞」），產生配子，配子又融合（「孢子生殖」）成二倍體合子，合子生長成「孢子囊」。然後，孢子囊進行減數分裂，形成傳播感染嘅孢子蟲。[63][150]

喺植物黏菌入面，二倍體「初級游動孢子」進入寄主，包囊，並作為單核「原生質體」或者「原質團」穿透細胞。喺細胞內部，原生質體生長成多核孢子囊，然後分裂成「次級游動孢子」，感染更多細胞。呢啲細胞繁殖成厚壁休眠孢子，休眠孢子開始減數分裂並分裂成雙核休眠孢子；一個細胞核消失，孢子孵化為初級游動孢子。[159]

一啲原生生物病原體喺好多唔同嘅生物（佢哋充當次要或者中間寄主）入面進行無性繁殖，但係淨係可以喺主要或者最終寄主入面進行有性繁殖（例如，弓形蟲 喺 貓科動物，好似家貓入面）。[160] 其他一啲，好似 利什曼原蟲屬，能夠喺次要嘅媒介中進行配子結合。[161] 喺頂複門入面，有性繁殖對於寄生蟲傳播嚟講係強制性嘅。[162]

即使進行有性繁殖，都唔清楚病原性原生生物唔同品系之間嘅基因交流有幾頻繁，因為大多數種群可能係克隆系，好少同佢哋物種嘅其他成員交換基因。[163]

原生生物對於全球現代生態系統嚟講係唔可以缺少嘅。[164] 喺 地球歷史 嘅大部分時間入面，佢哋亦都係所有生態系統嘅唯一真核生物成分，呢個令佢哋進化出廣泛嘅功能多樣性，解釋咗佢哋嘅關鍵生態學意義。佢哋作為初級生產者、多個營養級嘅中間體、關鍵嘅調節寄生生物或者寄生蜂，以及各種共生關係中嘅夥伴，都係必不可少嘅。[34]

原生生物喺幾乎所有棲息地都豐富而且多樣化。佢哋貢獻咗四十億噸 (Gt) 地球生物量——係動物嘅兩倍（2 Gt），但係少過全球生物量嘅 1%。加埋一齊，原生生物、動物、古菌 (7 Gt) 同真菌 (12 Gt) 加埋唔夠全球生物量嘅 10%，植物 (450 Gt) 同細菌 (70 Gt) 佔據主導地位。[165] 原生生物多樣性，正如通過 環境DNA 調查檢測到嘅噉，喺每個採樣環境中都好廣泛，但係大多數都未被描述。[166] 最豐富嘅原生生物群落出現喺土壤入面，其次係海洋棲息地，最後係淡水棲息地，大多數都係作為浮游生物嘅一部分。[167] 淡水原生生物群落嘅特點係具有更高嘅「β 多樣性」（即係樣本之間高度異質）。高多樣性可能係水文動態從唔同棲息地通過極端洪水度招募生物嘅結果。[168] 土壤棲息原生生物群落喺生態學上係最豐富嘅，可能係因為 沉積物 中水嘅分佈複雜且高度動態，呢個產生咗極其多樣化嘅環境條件。不斷變化嘅環境一次只促進群落嘅一部分嘅活動，而其餘部分保持唔活躍；呢種現象促進咗原核生物同原生生物嘅高度微生物多樣性。[167]


吞噬營養原生生物係所有生態系統中最多樣化嘅功能群體，主要由 Cercozoa（喺淡水同土壤中佔主導地位）、放射蟲（喺海洋中佔主導地位）、非光合作用 不等鞭毛類（喺土壤中嘅豐度高過喺海洋中）同 纖毛蟲 代表。[167]

同常見嘅浮游植物同動物浮游生物嘅劃分相反，海洋浮游生物嘅大部分都係由 混合營養 原生生物組成嘅，佢哋構成咗一個俾人嚴重低估咗嘅重要性同豐度（約莫佔所有海洋 環境DNA 序列嘅 12%）。混合營養生物嘅存在多樣，因為 季節 性豐度[174] 以及取決於佢哋特定嘅混合營養類型。構成型混合營養生物幾乎存在於海洋條件嘅所有範圍內，從富營養化嘅淺水棲息地到貧營養嘅亞熱帶水域，但主要喺 有光層 中佔據主導地位，佢哋佔咗細菌捕食嘅大多數。佢哋亦都係 有害藻華 嘅成因。質體型同通才非構成型混合營養生物具有相似嘅生物地理分佈同低豐度，主要喺富營養化嘅近岸水域度搵到，通才 纖毛蟲 喺有光層中嘅纖毛蟲群落中佔據咗一半以上嘅主導地位。最後，內共生混合營養生物係迄今為止最廣泛同最豐富嘅非構成型，佔咗所有混合營養序列嘅 90% 以上（大多數係 放射蟲）。[140][139]

土壤食物網嘅圖，考慮到原生生物作為唔單止係食菌生物，重有食真菌生物同雜食生物嘅多種角色。[175] 箭頭表示養分嘅流動。

喺土壤嘅 食物網 入面，原生生物係 細菌 同 真菌 嘅主要消費者，細菌同真菌係養分流向更高營養級嘅兩條主要途徑。[176] 變形鞭毛蟲，好似 Glissomonadida 同 Cercomonadida 係土壤原生生物中最豐富嘅：佢哋同時具有鞭毛同偽足，形態多樣性好適合喺土壤顆粒之間覓食。有殼變形蟲亦都適應咗土壤環境，因為佢哋嘅外殼可以抵抗乾燥。[173] 作為細菌食草動物，佢哋喺食物網中發揮住重要作用：佢哋以 氨 嘅形式排出 氮，令到植物同其他微生物都可以利用到。[176] 傳統上，原生生物俾人認為主要係食菌性嘅，因為培養技術存在偏差，但係好多（例如，Vampyrellida、Cercomonadida、裸變形蟲、有殼變形蟲、細小鞭毛蟲）都係雜食性嘅，佢哋以各種各樣嘅土壤真核生物做嘢食，包括真菌，甚至一啲動物，好似線蟲。食菌性同食真菌性原生生物嘅生物量差唔多。[175]

比奇利認為呢個界太多多系群，並拒絕將細菌納入其中。佢將原生生物界分割成「原生動物」（淨係有核、單細胞、類似動物嘅生物），而細菌同「原生植物」就係一個獨立嘅分組。呢個加強咗德國科學家 卡爾·西奧多·馮·西博爾德 嘅舊二分法「原生動物」/「原生植物」，而喺個世紀之交，德國博物學家向全球科學界確立咗呢個觀點。但係，英國生物學家 C. 克利福德·多貝爾 喺 1911 年提請注意，原生生物嘅功能同動物同植物細胞組織嘅功能非常唔同，並且重視原生生物作為一個具有唔同組織嘅群體，佢將呢種組織叫做「無細胞性」，從而擺脫咗德國細胞理論嘅教條。佢創造咗 原生生物學 呢個術語，並鞏固咗佢作為一個獨立於 動物學 同 植物學 嘅研究分支。[25]

喺 1938 年，美國生物學家 赫伯特·科普蘭 復活咗霍格嘅標籤，佢認為海克爾嘅術語 原生生物 包括無核微生物，例如細菌，而術語 原生生物總界 （意思係「第一批建立嘅生物」）就冇包括。喺佢嘅四界分類（原核生物界、原生生物總界、植物界、動物界）之下，原生生物同細菌最終分開，承認咗無核（原核生物）同有核（真核生物）生物之間嘅區別。為咗將原生生物同植物完全分開，佢遵循海克爾對真正動物嘅囊胚定義，並提出將 真正植物 定義為具有 葉綠素 a 同 b、胡蘿蔔素、葉黃素 同 澱粉 產生嘅植物。佢亦都係第一個意識到單細胞/多細胞二分法係無效嘅人。儘管如此，佢仍然將真菌同 紅藻、褐藻 同 原生動物 保留喺原生生物總界入面。[25][194] 呢個分類係惠特克之後對真菌、動物界、植物界 同原生生物界作為生命嘅四個界嘅定義嘅基礎。[195]

喺美國植物生態學家 羅伯特·惠特克 喺 1969 年發表嘅流行 五界 方案入面，原生生物界俾定義做真核生物「單細胞 或者單細胞-群體，而且唔形成 組織」嘅生物。正如原核生物/真核生物嘅劃分變得主流一樣，惠特克喺科普蘭系統嘅十年之後，[195] 承認咗生命喺原核生物原核生物界同真核生物界之間嘅根本劃分：動物界（攝食）、植物界（光合作用）、真菌界（吸收）同剩低嘅原生生物界。[196][197][25]

喺美國進化生物學家 琳·馬古利斯 嘅五界系統中，「原生生物」呢個術語保留俾 微觀生物，而更具包容性嘅原生生物總界（或者 原生生物總界生物）包括一啲大型多細胞真核生物，好似 海帶、紅藻 同 黏菌。[198] 一啲人將「原生生物」呢個術語同馬古利斯嘅「原生生物總界生物」互換使用，嚟涵蓋單細胞同多細胞真核生物，包括嗰啲形成特殊組織，但係唔符合任何其他傳統界別嘅生物。[199]

  1. 1 2 原生生物同其他三個真核生物界之間嘅區別一直都好難確定。喺歷史上，異養原生生物，叫做原生動物，俾人認為係動物界嘅一部分，而自養原生生物，叫做藻類，就俾人當做植物界嘅一部分嚟研究。即使喺建立咗一個獨立嘅原生生物界之後，一啲微細嘅動物（黏體動物）[3] 同「較低等」嘅真菌（即係Aphelida、Rozellida 同 微孢子蟲，統稱為 Opisthosporidia) 都俾人當做原生生物嚟研究，[4][5][6] 而且一啲藻類（尤其係紅藻同綠藻）仍然俾人歸類做植物。[7] 根據目前嘅共識，「原生生物」呢個標籤明確噉排除咗動物、胚胎植物（陸地植物）——意思係所有藻類都屬於呢個標籤——同埋所有真菌。後鞭毛菌門俾人認為係較大嘅真菌界嘅一部分，即使佢哋通常都俾原生生物學家同真菌學家共同研究。[8][9][10]
  2. ↑ 從生物學嘅角度嚟講，「纖毛」同「真核鞭毛」呢啲術語可以互換使用。但係，佢哋嘅用法取決於作者：一啲人更鍾意將纖毛保留俾較短嘅附屬物，而將鞭毛保留俾較長嘅附屬物，而另一些人更鍾意將纖毛用於真核生物，而將鞭毛用於原核生物。「軸絲足」呢個術語俾人提出嚟統一呢兩個概念，因為佢特指喺兩者中都搵到嘅同源微管結構，但係喺原核生物鞭毛中就搵唔到。[21][22][23]
  3. ↑ 一份關於原生生物多樣性嘅 2007 年報告包含一個表格，列出咗原生生物同真菌群體嘅已描述物種數量。列出嘅物種總數，唔包括真菌，係 76,144。[33]
  4. 1 2 門 Percolozoa 通常更出名嘅係異葉足蟲綱。[8][40] 但係，喺最嚴格嘅意義上，異葉足蟲綱僅指呢個門內嘅一個綱，包含 Acrasida 同 Schizopyrenida 目。名稱 Percolozoa 包括呢啲同其他相關嘅單細胞原生生物，唔單止係「真正」嘅異葉足蟲。[6]
  5. ↑ 「混合營養生物」同「混合浮游生物」呢啲術語幾乎專門指嘅係進行光合作用同吞噬作用嘅原生生物（光合-吞噬營養生物）。滲透營養一直都存在，但係冇俾人考慮在內。因此，「純粹」嘅光合營養生物（冇吞噬作用能力）同「純粹」嘅吞噬營養生物（冇光合作用能力）喺技術上都係混合營養生物，因為佢哋天生具有滲透營養嘅能力，但係通常唔會喺呢個意義上報告。[138]
  6. ↑ 卡爾·林奈 直到 1758 年第十版嘅 自然系統 先至提及到任何一個原生生物屬，喺嗰度， 團藻 俾人記錄咗。[185]
  7. ↑ 喺 2015 年，卡瓦利耶-史密斯最初嘅六界模型俾修訂做 七界模型，喺納入 古菌 之後。[207]

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參考書目

通用

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Margulis, L.、J.O. Corliss、M. Melkonian、D.J. Chapman. 原生生物總界手冊. Jones and Bartlett Publishers, Boston, 1990.

Margulis, L.、K.V. Schwartz. 五界：地球生命門嘅圖解指南, 第 3 版。 New York: W.H. Freeman, 1998.

Margulis, L.、L. Olendzenski、H.I. McKhann. 原生生物總界圖解詞彙表, 1993.

Margulis, L.、M.J. Chapman. 界同域：地球生命門嘅圖解指南. Amsterdam: Academic Press/Elsevier, 2009.

Schaechter, M. 真核微生物. Amsterdam, Academic Press, 2012.

生理學、生態學同古生物學

Fontaneto, D. 微觀生物嘅生物地理學。細嘢係咪乜嘢地方都有？ Cambridge University Press, Cambridge, 2011.

Moore, R. C.，同其他編輯。 無脊椎動物古生物學專論。 Protista，B 部分（第 1 卷[失咗效嘅鏈]，Charophyta，第 2 卷，金黃藻目，顆石藻目，Charophyta，矽藻目同火藻目），C 部分（肉足蟲，主要係「有殼變形蟲」同有孔蟲目）同 D 部分[失咗效嘅鏈] （主要係放射蟲同 Tintinnina）。 Boulder, Colorado: Geological Society of America; & Lawrence, Kansas: University of Kansas Press.