【食品課件】真核微生物的形態(tài)結構和功能95

1、Chapter 2Morphology，structure and physiological function of eukaryotesDefinition： eukaryotic cells are compartmentalized by membran-es .The cell contains several different types of membrane bou- nd organelle in which different biochemical and physiological processes can occur in a regulated way. Mem

2、branes also transport information， metabolic intermediats and end-products from the site of biosynthesis to the site of use.Eukaryotes is a big group of living organisms made up of eukaryotic membrane bound nucleus and several organelles such as mitochondrion divided by mitosis.真核生物是一大類細胞核具有核膜，能進行有絲

3、分裂，細胞質中存在線粒體或同時存在葉綠體等多種細胞器的生物. eukaryotic micro-organisms is a group of Microorganisms that made up of eukaryotic cells include Fungi、microalgae、and protozoa真核細胞真核生物 真核微生物peroxisomesmitochondrioncytosolchloroplastSmooth endoplasmic reticulumStructure of a typical plant cellCytosol細胞液Plasma membraneS

4、ecretory vesiclesStructure of a typical animal cellmembraneCell wall mitochondrionVesicals (泡囊) Dictyosomes (分散高爾基體)Hyphal structuresEndoplasmic reticlumribosomesmembranevacuoleCell wallmitochondrionNucleus Endoplasmic reticulumYeast structuresYeast structures 真核微生物主要包括菌物界(Mycetalia或廣義的“Fungi)中的真菌(E

5、umycota或狹義的“Fungi，即True Fungi)、粘菌(Myxomycota或Fungilike Protozoa)、假菌(Chromista或Pseudofungi)，植物界(Plantae)中的顯微藻類(Algae)和動物界(Animalia)中的原生動物(Protozoa)Major groups of eukaryotic microorganisms 真核微生物的主要類群 植物界 plantae：顯微藻類 algae 動物界animalia：原生動物 protozoa 真核微生物 黏菌myxomycota 假菌pseudofungi 菌物界mycetalia 單細胞真菌u

6、nicellular fungi 真菌 eumycota 絲狀真菌filamentous 大型子實體 蕈菌 mushroomMajor groups of eukaryotic microorganisms Eukaryotic microorganisms“菌物界這個名詞是我國學者裘維蕃等于1990年提出的，并已得到學術界的一定支持，這是指與動物界、植物界相并列的一大群無葉綠素、依靠細胞外表吸收有機養(yǎng)料、細胞壁一般含有幾丁質的真核微生物。一般包括真菌、粘菌和假菌(卵菌等)3類。真菌是最重要的真核微生物，故是本章的重點，它們的特點是：無葉綠素，不能進行光合作用；一般具有興旺的菌絲體；細胞壁多數(shù)

7、含幾丁質；營養(yǎng)方式為異養(yǎng)吸收型；以產(chǎn)生大量無性和(或)有性孢子的方式進行繁殖；陸生性較強Fungi are filamentous, non-photosynthetic, eukaryotic microorganisms that have a heterotrophic nutrition . Their basic cellular unit is described as a hypha . This is a tubular cell which is surrounded by a rigid, chitin-containing cell wall. The hypha ext

8、ends by tip growth, and multiplies by branching, creating a fine network called a mycelium. Hyphae contain nuclei, mitochondria, ribosomes, Golgi and membrane-bound vesicles within a plasma-membrane bound cytoplasm . The subcellular structures are supported and organized by microtubules and endoplas

9、mic reticulum. The cytoplasmic contents of the hypha tend to be concentrated towards the growing tip. Older parts of the hypha are heavily vacuolated and may be separated from the younger areas by cross walls called septae. Not all fungi are multicellular, some are unicellular and are termed yeasts.

10、 These grow by binary fission or budding, creating new individuals from the parent cell.Mould Mold Definition Mold are filamentous, nonphotosynthetic, eukaryotic microorganisms that have a heterotrophic nutritionfilamentous fungi。 Morphology and structureHypha and mycelium菌絲與菌絲體 lower funginon-septa

11、te myceliums higher fungiseptate mycelium with elaborate、 perforate septa The hypha extends by tip growth, and multiplies by branching, creating a fine network called a mycelium. plural myceliaMycelia can be divided into vegatative mycelium and aerial mycelium營養(yǎng)菌絲體和氣生菌絲體Basic unithyphapl。 hyphae Thi

12、s is a tubular cell which is surrounded by a rigid, chitin-containing cell wall. Hyphae contain nuclei, mitochondria, ribosomes, Golgi and membrane-bound vesicles within a plasma-membrane bound cytoplasm membraneCell wall mitochondrionVesicals Dictyosomes (分散高爾基體)Hyphal structuresSpecialized morphol

13、ogy of vegetative mycelium營養(yǎng)菌絲體的特化形態(tài)Rhizoid 假根Rhizopus Stolon 匍匐菌絲Mucorales 、Rhizopus Haustorium 吸器Adhesive cell 附著胞Adhesive branch 附著枝Sclerotium 菌核Rhizomorph ，funiculus菌索 ring and net 菌環(huán)和菌網(wǎng)1)假根(rhizoid) 是Rhizopus(根霉屬)等低等真菌匍匐菌絲與固體基質接觸處分化出來的根狀結構，具有固著和吸取養(yǎng)料等功能 (2)匍匐菌絲(stolon)又稱匍匐枝。毛霉目(Mueorales)真菌在固體基質

14、上常形成與外表平行、具有延伸功能的菌絲，稱匍匐菌絲。最典型的可在Rhizopus中見：在固體基質外表的營養(yǎng)菌絲分化為匍匐菌絲，在其上每隔一段距離可長出伸人基質的假根 根霉的形態(tài)和構造和伸向空間的孢囊梗，隨著匍匐菌絲的延伸，不斷形成新的假根和孢囊梗，這類真菌會隨基質的存在而向四處快速蔓延，根本就不會形成像在其他真菌中常見的那樣有固定大小和形態(tài)的菌落。sporangium(3)吸器(haustorium) 由幾類專性寄生的真菌如銹菌目(Uredinales)、霜霉目(Peronosporales)和白粉菌目(Erysiphales)等的一些種所產(chǎn)生。吸器是一種只在宿主細胞間隙間蔓延的營養(yǎng)菌絲上分化

15、出來的短枝，它可侵入細胞內形成指狀、球狀或絲狀的構造，用以吸取宿主細胞內的養(yǎng)料而不使其致死。(4)附著胞(adhesive cell) 許多寄生于植物的真菌在其芽管或老菌絲頂端會發(fā)產(chǎn) 膨大，分泌粘狀物，借以牢固地粘附在宿主的外表，此即附著胞。在其上再形成針狀感染菌絲，以侵入宿主的角質表皮而吸取養(yǎng)料。 (5)附著枝(adhesive branch) 假設干寄生真菌由菌絲細胞生出12個細胞的短枝，將菌絲附著于宿主體上，6菌核sclerotium) 是一種形狀、大小不一的休眠菌絲組織，在不良外界條件下：可保存數(shù)年生命力。菌核形狀有大有小，大的如茯苓(大如小孩頭)，小的如油菜菌核(形如鼠糞。菌核的外層

16、色深、堅硬、內層疏松，大多呈白色。7菌索一般由傘菌等產(chǎn)生，為白色根狀菌絲組織，功能為促進菌體蔓延和抵御不良環(huán)境。通?？稍诟嗟臉淦は潞偷叵掳l(fā)現(xiàn)。 (8)菌環(huán)(loop )和菌網(wǎng)( net) 捕蟲菌目(zoopasdes)和一些半知菌的菌絲常會分化成圈環(huán)或網(wǎng)狀的特化菌絲組織，用以捕捉線蟲或其他微小動物，然后進一步從這類環(huán)或網(wǎng)上生出菌絲侵入線蟲等體內，吸收養(yǎng)料9子實體fruiting body，sporocarp， fructificationColonial growthHyphal tip growth allows fungi to extend into new regions from

17、a point source or inoculum(接種物). Older parts of the hyphae are often emptied of contents as the cytoplasm is taken forwards with the growing tip. (老的菌絲通常缺乏內含物，因為細胞質流向生長點)This creates the radiating colonial pattern seen on agar plates , in ringworm infections of skin and fairy rings in grass lawns. O

18、nly hyphal tips contribute to extension growth. However older hyphae can grow aerially or differentiate to produce sporing structures只有菌絲頂端對延伸生長起作用，而老菌絲可以進行氣生生長或分化出產(chǎn)孢子的結構Mycelium in agarAgar Surface myceliumKinetics of growth Fungal growth in a given medium follows the growth phases of lag, accelera

19、tion, exponential, linear, retardation, stationary and decline延遲期、加速期、指數(shù)期、直線期、減速期、穩(wěn)定期、衰亡期 .Exponential growth occurs only for a brief period as hyphae branches initiated, and then the new hypha extends at a linear rate into uncolonized regions of substrate. Life cycles All fungi are characterized by

20、 having a period of vegetative growth where their biomass increases. The length of time and the amount of biomass needed before sporulation can occur varies. Almost all fungi reproduce by the production of spores, but a few have lost all sporing structures and are referred to as mycelia sterilia(不育菌

21、絲體). Different types of spore are produced in different parts of the life cycle.Reproduction in fungiAll fungi undergo a period of vegetative growth where their mycelium exploits a substrate. This stage is followed by asexual and sexual reproduction. There are two conflicting requirements fungi have

22、 for their spores. Spores must allow fungi to spread, but they must also allow them to survive adverse conditions. These requirements are met by different types of spores. Small, light spores are carried furthest from parent mycelium in air and these are the dispersal spores釋放型孢子. They are usually t

23、he products of asexual sporulation, the sporangiospores and the conidiospores, and so spread genetically identical individuals as widely as possible. Genetic diversity is maintained by sexual reproduction, and the spore products are often large resting spores休眠孢子 that withstand adverse conditions bu

24、t remain close to their site of formation. Spores therefore vary greatly is size, shape and ornamentation, and this variation reflects specialization of purpose. Fungal spores：spores allow fungi to spread, to maintain genetic diversity and to survive adverse conditionsReproduction in fungi spore dis

25、charge ：Spores that have a dispersal function can be released from their parent mycelium by active or passive mechanisms . As many spores are wind dispersed, they are produced in dry friable masses which are passively discharged by wind. Other spores are passively discharged by water droplets splash

26、ing spores away from parent mycelium. Spores may be discharged from parent mycelium by passive or active means. Passive mechanisms include using wind and water as dispersants;active mechanisms use explosive principles.Fungal spore discharge Spores in the atmosphereair-borne fungal spores can be carr

27、ied great distances。Their presence in the air can have impact on human health as they can cause allergic rhinitus hay feverand asthma。 Many plant diseases that cause great economic losses are air borne。can affect human, animal and plant health. They can cause allergies and spread plant disease.Air s

28、poraFungi reproduce by the formation of sexual or asexual spores Asexual reproductionfission ArthrosporeBudding blastosporeOther asexual spore Zoospore Chlamydospore Sporangiospore conidiospora Sexual reproduction Oospore Zygospore Ascospore Basidiospore節(jié)孢子(arthrospore)某些真菌生長到一定階段，菌絲中間形成許多隔膜，接著從隔膜處斷

29、裂成許多竹節(jié)似的無性孢子，稱為節(jié)孢子，也稱為粉孢子。最典型的例于是白地霉(Geotrichumcandidum).arthrosporeBlastospore (芽孢子) 這和酵母菌的出芽一樣，它是由母細胞生芽而形成的。當芽長到正常大小時，脫離母細胞，或仍連在母細胞上。如玉蜀黍黑粉菌(Ustilago maydis)能產(chǎn)生芽孢子。某些毛霉或根霉在液體培養(yǎng)基中形成的被稱為酵母型細胞，也屬芽孢子。 又稱厚壁孢子或厚膜孢子。某些真菌生長到定階段，在菌絲的頂端或中間有局部細胞的細胞質密集在一起，變圓，然后在其四周生出厚壁，或原細胞壁加厚，形成圓形、紡錘形的無性休眠體，來抵抗外界不良的環(huán)境條件。例如總狀

30、毛霉(Mucor racemosus)往往在菌絲中間形成許多厚垣孢子schlamydospore厚垣孢子 無性繁殖產(chǎn)生的孢子在孢子囊(Sporange)內,孢子囊一般生在氣生菌絲的頂端或生在孢囊梗(sporangiophore)的頂端。在形成孢子囊前，首先有多核的原生質密集于此處，使其膨大，并在下方生出橫隔，形成圓形的囊狀物，然后其中原生質體割裂成許多小塊，每塊發(fā)育成為個孢子囊孢子。因而每一個孢子囊所含有的孢子的數(shù)般都相當多。 就孢子囊孢子而言,般有二類,一種具有根或23根鞭毛，能夠游動，所以稱游動孢子(zoospore)。例如腐霉(即thium)。另一種無鞭毛，不能游動，又稱靜止擔于。spo

31、rangiospore (孢子囊孢子)游動孢子 conidium (分生孢子) 分生孢子的形成方法 其形成方式有兩種，一種是在分生孢子梗的頂端突出，發(fā)育成第一個孢子。梗再伸長形成第二個孢子，如此重復形成一串孢子。這樣形成的分生孢子，頂端的最老。曲霉屬和青霉屆的分生孢子形成，即屬于此類型。 另一種是在第個分生孢子形成時，柄的長度已到達最高，由第一個分生孢子頂端生長出第二個分生孢子，如此重復形成一串孢子。這樣形成的分生孢子，下部的最老。如枝孢霉屬(Cladosporium)的分生孢子形成即屬于此類型。Plastic kitchen cutting board surface - rod bacte

32、ria in a smooth matrix; filamentous bacteria ( 絲狀菌 ) -smooth oval cells; fungal hyphae ( 真菌菌絲 ) with textured spores ( 結構孢子 ) (x 2,270)conidiumZoospore游動孢子Asexual spore， usually produced by fungi living in water .Motile zoospores in motile sporangia differentiated from swelled hyphal tips have a sin

33、gle posterior flagellum。The structure of the flagellum is 9+2 ，some has two flagella. 霉菌的有性繁殖是經(jīng)過不同性別的細胞結合(質配和核配)后，產(chǎn)生一定形態(tài)的孢子來實現(xiàn)的，這種孢子稱為有性孢子。繁殖過程可分為三個階段：第個階段為質配；第二個階段為核配，產(chǎn)生二倍體的核；第三個階段是減數(shù)分裂，恢復核的單倍體狀態(tài)。大多數(shù)真菌菌體是單倍體的。有性孢子通常有以下幾種：Sexual reproduction 菌絲分為雄器(antheridium)和藏卵器(oogonium)。藏卵器中有一個或數(shù)個卵球(oosphere)。當

34、雄器和藏卵器相配時，雄器中細胞質與細胞核，通過受精管而進入藏卵器，與卵球結合形成卵孢子。圖所示是德巴利腐霉的卵孢子的形成。 oospore (卵孢子) b同宗結合a異宗結合根霉的接合孢子Zygospore接合孢子ascospore 子囊孢子形成子囊孢子是子囊菌的主要特征。子囊中孢子數(shù)目通常為18個，或為2n。典型的子囊中有8個孢子。大多數(shù)霉菌的子囊是包裹在子囊果ascocarp中。子囊果有三種類型 子囊殼 perithecium 子囊盤apothecium 閉囊殼cleistothecium Sexual reproduction in this group occurs after soma

35、tic fusion of different mating-type mycelia. A transient diploid phase is rapidly followed by the formation of ascospores within sac-shaped asci differentiated from modified hyphal tips. In the initial stages of ascal development hooked hyphal tips form, called croziers 產(chǎn)囊絲鉤or shepherds crooks牧羊杖 be

36、cause of their shape. They have distinctive septae at their base which insure that two different mating-type nuclei are maintained in the terminal cell. Formation of the septae is coordinated with nuclear division . In yeasts all these events occur within one cell, after fusion of two mating-type ce

37、lls, the whole cell being converted into an ascus.The formation of ascosporesSexual reproduction in the ascomycetes In more complex Ascomycetes many asci form together, creating a fertile tissue called a hymenium叫子實層的能育組織. In some groups the hymeniumcan be supported or even enclosed by large amounts

38、 of vegetative mycelium. The whole structure is called a fruit body or sporocarp子囊果 and is used as a major taxonomic feature . They can become large enough to be seen with the naked eye. Flask-shaped sexual reproductive bodies are called perithecia, cup-shaped bodies are called apothecia and closed

39、bodies are called cleistothecia. These structures have evolved to protect the asci in spore dispersal， but the hymenium itself is unafected by the presence of water。The formation of sporocarp and its formsCup-shaped apotheciumFlask-shaped peritheciumClosed cleistotheciumAsci Asci Ascus containing as

40、cosporesBasidiomycete fungi rarely reproduce asexually. Sexual reproduction is by the formation of basidiospores on the gills or pores of large fruit group of fungi are characterized by the most complex and large structures found in the fungi. They are also distinctive in that they very rarely produ

41、ce asexual spores. Much of the life cycle is spent as vegetative mycelium, exploiting complex substrates. A preliminary requisite for the onset of sexual reproduction is the acquisition of two mating types of nuclei by the fusion of compatible hyphae. Single representatives of the two mating-type nu

42、clei are held within every hyphal compartment for extended periods of time. This is termed a dikaryotic state雙核期, and its maintenance requires elaborate septum formation during growth and nuclear division .Basidiospore Onset of sexual-spore formation is triggered by environmental conditions and begi

43、ns with the formation of a fruit body primordium子實體原基. Dikaryotic mycelium雙核菌絲體 expands and differentiates to form the large fruit bodies we recognize as mushrooms and toadstools毒蕈. Diploid formation and meiosis occur within a modified hyphal tip called a basidium 。Basidium formation 核融合Nuclear fusi

44、on減數(shù)分裂meiosis 擔孢子梗sterigmataBasidiospores擔孢子Basidium擔子Basidium and basidiospore formationFour spores are budded from the basidium. Basidia form together to create a hymenium which is highly sensitive to the presence of free water. The hym-enium is distributed over sterile, dikaryotic supporting tiss

45、ues which protect it from rain. The hymenium can be exposed on gills or pores beneath the fruit body, seen in the toadstools and bracket fungi, or enclosed within chambers as in the puffballs and truffles Formation of basidiosporepuffball 馬勃Bracket檐狀傘 Toadstool毒蘑菇Hymenenium子實層Structure of sexual spo

46、rocarps in the basidiomycetesbasidiospore擔孢子擔孢子是擔子菌特有的特征。它是一種外生孢子，經(jīng)過兩性細胞核配合后產(chǎn)生。因為它生在擔子上，所以稱為擔孢子。典型的擔子菌的擔子上有四個擔孢子。各種擔孢子的形狀及外表特征Food related molds1. Aspergillus 曲霉屬 2. penicillium青霉屬 3. Rhizopus 根霉屬4. sporotrichum (側孢霉屬)5. Thamnidium(枝霉屬)6. Alternaria交鏈孢屬7.Botrytis 葡萄孢霉木霉屬紅曲屬赤霉屬 11.Cordyceps蟲草屬地霉屬長蠕孢霉

47、food related MouldsAspergillus曲霉屬 分生孢子 初生小梗孢子穗 小梗 次生小梗 頂囊 足細胞 Conidia primary sterigmaconidial head sterigme secondary sterigma Vesicle foot cellMorphology of Conidial head is the basis for species identification孢子穗的形態(tài)是菌種鑒定的依據(jù)Aspergillus Relationship with foodbeneficialFermentation industry soy sauc

48、e釀醬 vinegar制醋曲 fermented bean curd腐乳 alcohol beverage釀酒Food processing organic acid有機酸 enzyme 酶制劑 淀粉酶 蛋白酶 果膠酶等Relationship with foodharmful Usually distributed on the surface of following foodstuff：Cakes、fruits、vegetables、meat、 grain and other organic objects。Causing following consequences：Go mouldy

49、；become mildewedProduce or secrete carcinogenic substanceblastomogen致癌物質eg 。 aflatoxin黃曲霉毒素Food related moldspenicillium青霉屬 十分接近于曲霉，在自然界分布很廣，長生長在腐爛的柑橘皮上，呈青綠色，不少種類引起食品變質，但也用來生產(chǎn)青霉素和有機酸等。 青霉菌菌絲與曲霉相似，但無足細胞，孢子穗結構也不同。其分生孢子梗頂端不膨大，而是經(jīng)過屢次分枝產(chǎn)生幾輪對稱或不對稱的小梗，然后在小梗的頂端產(chǎn)生成串的分生孢子。青霉菌的孢子穗形狀似掃帚狀。分生孢子球形、橢圓形或短柱形，一般呈藍綠色。

50、Plastic kitchen cutting board surface - rod bacteria in a smooth matrix; filamentous bacteria ( 絲狀菌 ) -smooth oval cells; fungal hyphae ( 真菌菌絲 ) with textured spores ( 結構孢子 ) (x 2,270)根霉在自然界分布廣泛，經(jīng)常出現(xiàn)在淀粉質食品上，引起糧食、食品腐爛、腐敗。I工業(yè)上常利用根霉生產(chǎn)糖化酶，甜酒曲中主要菌種也是根霉。此外，根霉也可以用來生產(chǎn)有機酸等其它物質。Food related moldsRhizopus 根霉屬R

51、hizoid假根Stolen匍匐絲 sporangeUaually blacksporangiophore 菌絲分隔分生孢子梗分枝，頂端產(chǎn)生分生孢子。分生孢子單細胞，梨形。菌落呈奶油色澤，時間長后，為枯燥粉末狀。已經(jīng)報告該菌能在OC以下生長，常在冷藏肉中形成白色斑點。 food related moldssporotrichum (側孢霉屬)SCarnis肉色側孢霉Conidium分生孢子Conidiophore分生孢子梗Cause the deterioration of vegetables and fruitconidiumconidiophore形狀大小不定，數(shù)個連接成鏈，暗褐色，有縱

52、橫隔膜，呈壁磚狀。大多數(shù)不分支，較短，單生或成簇，暗褐色。Food related moldsAlternaria交鏈孢屬septate mycelium Food related moldsBotrytis 葡萄孢霉conidiophoreconidium分生孢子梗頂端形成樹枝狀分支，分支頂端細胞常常膨大，在短的小梗上著生分生孢子，如一串葡萄。卵圓形，無色或暗褐色。常產(chǎn)生外形不規(guī)那么的黑色菌核sclerotium。分布在許多植物和植物食品上，形成一層“灰色霉，引起水果、蔬菜的腐敗。Food related moldsTrichoderma木霉屬Spherical or ellipticalo

53、val，smmoth or rough，yellow-green conidiaColorless hypha，with septa and branches ，produce chlamydosporesBottle-like sterigme瓶狀小梗Spore head gatheredconglutenated by mucusslimeFood related moldsTrichoderma木霉屬Colony grows rapidly，like cotton fibre棉絮狀，white at first，later become green。Spore-producing are

54、a takes onhas concentric circlesSone species has cellulolytic activity，therefore can be used to produce cellulase to manufacture sugars with cellular waste下腳，starch processing and food processing Moldy grain，fruit and vegetablesMildew and rot of cellular material。Food related moldsMonascus紅曲屬Can sec

55、rete red pigmentThe source of natural red pigment in the food processing，for instance red riceang-kakused in the processing of red fermented bean curd，bevarage and meat is prepared from Monascus。Type culture that commonly used：M. purpureus紫紅曲Food related moldsMonascus紅曲屬Colony features： membranous c

56、olony covers the whole malt extract plate，mycelium is white at first，later becomes red，red-purple，the water-soluble pigments are secreted into the medium。 conidiumFood related moldsMonascus紅曲屬13 initiate cleistothecium45 premature cleistothecium 6 mature cleistothecium cleistothecium of 紫紅曲的閉囊殼Food

57、related moldsGibberella赤霉屬 reproduction scarcely produce Sexual generatin gamobium ，sexual progeny，sexual spore is ascospore in spherical ，smooth and blue perithecium子囊殼。 Asexual reproduction produce two types of conidiumSmall type：single cellular，spherical，elliptical to long column，colorless or oth

58、er colorsLarge type：multicellular with septa， Sickle-like or long column，Colorless or other colorsFood related moldsGibberella赤霉屬 Gibberellin or abbeymycin 赤霉素 Metabolic product of Gibberella，is a kind of hormone with the following function：1.Stimulate plant growth2.Break the dormancy of seed and tu

59、ber 3.Increase the yield of vegetables especially leafy vegetablesFood related moldsCordyceps蟲草屬Formation： The insect body parasitized by the fungi was full of myceliumsclerotium 菌核and mummified， stem-head like or rod like stroma plural stromata 子座grows out from its front end。Inchworm尺蠖 Stroma grass

60、usually single， scarcely 2 or 3，411cm in length。peritheciumAscusasciascosporeCordyceps sinesis冬蟲夏草Food related moldsgeotrichum地霉屬Reproduction by fission，septate hypha，hypha may break into arthrospore，without blastospore。 Always existed in sauerkrautkimchi，pickles泡菜， fecund，organic fertilizer，rotten