Phakathi kwezinambuzane, izindlela eziyinkimbinkimbi kakhulu zokuziphatha ziyabonakala ezinambuzaneni zomphakathi. Inhlangano yemiphakathi yayo isuselwa ebudlelwaneni nasekusebenzisaneni phakathi kwamalungu ayo ahlala koloni eyodwa nokuhlala esidlekeni sohlobo oluthile noma kolunye. Izinambuzane zomphakathi zangempela, okuthiwa izinambuzane ze-eusocial, zingama-oda amabili. Konke lokhu kungama-termites (Isoptera) nabamele abakhethekile beHymenoptera. zonke izintuthwane neminye iminyovu ehlelwe kahle kakhulu nezinyosi.
Ukuziphatha kwe-Eusocial kubonakaliswa yizici ezintathu eziphambili.
- koloni lwale zinhlobo, abantu bahlangana ukunakekela izingane,
- imisebenzi koloni isatshalaliswa ngamaqembu akhethekile abantu,
- imijikelezo yempilo yabantu okungenani okungenani yezizukulwane ezimbili, ukuze isizukulwane esincane sichitha ingxenye yezimpilo zabo kanye nesizukulwane somzali.
Kwamazinga wangaphambi kokuhlalisana (presocial) kwenhlangano yezinambuzane, munye noma ezimbili kulezi zici ezintathu ezinesimo.
Amakoloni ezinambuzane ze-Eusocial nawo abonakala ngezinga eliphakeme lokuzala. Inani labantu kule koloni lisuka ngaphansi kwezigidi eziyi-100 liye kwizigidi ezinhlobonhlobo ezihlukene kanye namaqembu ezinambuzane (Umd. 6.25, 6.26). Ekholoni lenye yezinhlobo zezintuthwane zase-Afrika kungafinyelela abantu abayizigidi ezingama-22. Ezinye izintuthwane ezihleleke kakhulu, ngokwesibonelo I-Myrmica rubra, koloni kukhona isibeletho esisodwa ("indlovukazi"), esihlala iminyaka eminingana, sibeka amaqanda futhi sinikeze inzalo kulo lonke leli koloni, futhi zonke izinsikazi ezingazaleli ziyinika ukudla. Abantu abangazaleli benza imisebenzi eminingi koloni.
Imvamisa amelelwa izinhlobo ezahlukahlukene ze-morphological, noma ama-castes. Abantu abancane kakhulu - izisebenzi, ezinkulu - amasotsha noma izisebenzi ezinkulu. Abasebenzi bazibandakanye ekuqoqeni okuphakelayo, ekudleni, ekunakekeleni inzalo. Amasosha avikela isidleke nezindawo zokugcina ukudla okungamanzi. Abantu abangazalanga baphila isikhathi esifushane, kanti isibeletho kufanele sibelethe amaqanda cishe ngokuqhubekayo ukuqinisekisa ukuthi ikoloni likhulu ngokwanele ngesikhathi sokuvuthwa. Ngaphezu kwalawa maqembu omphakathi, kukhona elinye iqembu eliqukethe abesilisa. Azenzi noma yimuphi umsebenzi, zinemisebenzi ethile yezenhlalo (ngokwesibonelo, enakekela abanye abantu) futhi zilinde indiza yokuya esikhathini ukuthi ifake izintokazi ezingazili. Izinsikazi zezintombi zikhula kusukela kuzalo lomkhiqizi wesifazane. Ngemuva kwenkathi yokuzalela, insikazi ngayinye isungula ikhole layo elisha, yakha isidleke futhi inakekele inzalo. Lapho ikoloni selivuthiwe, abantu abangazali banakekela inzalo neminye imisebenzi.
Abanduleli bokuziphendukela kwenqanaba le-eusocial yenhlangano yezinambuzane babe yimigqa emibili yokuthuthukiswa kobudlelwano bezenhlalo - i-parasocial futhi -phansi kwezenhlalo (Fig. 6.27). Ngendlela yokuphila yodwa (engeyona eyezenhlalo), akunakukhathalela inzalo, azikho izinkundla ezikhethekile zokuzalela kanye nokugqagqana kwezizukulwane ngezizukulwane ezilandelanayo. Ngenhlangano yenhlangano eyisimo sezinyosi zomndeni wakwaHalictidae, abantu abadala besizukulwane esifanayo, ngezinga elithile noma kwelinye, bayasizana. Izinga eliphansi lenhlangano yalolu hlobo ibizwa ngokuthi bomphakathi. Emiphakathini okulesi sigaba, abantu abadala bahlangana ukwakha isidleke, kepha inzalo ikhuliswa ngokwahlukana.
Ezingeni lokuhlangana kukhona izinhlobo eziningana zezinyosi ezivela emndenini weHalictidae. Cishe abantu besifazane abangama-50 bohlobo olufanayo bahlala esidlekeni esingaphansi komhlaba. Inyosi ngayinye ibamba imigudu yayo noma amaseli ayo aseceleni, lapho ibeka khona iqanda elilodwa, ibeka ukudla bese kuyivala. Ezingeni elilandelayo lomugqa we-parasocial - Quasi-social - Ukunakekela inzalo kungokwesimo somphakathi, kepha owesilisa ngamunye ubeka amaqanda ngesikhathi esithile. Kokulandelayo, hemisocial, kuvela umqulu wabasebenzi, onabantu abangazalanisi bekholoni. Ileveli elandelayo yenhlangano isivele ikhona okululaotholakala lapho isikhathi sokuphila kwesinye sezizukulwane se-hemisocial colony siba sikhulu kangangokuba izizukulwane ezimbili noma ngaphezulu zilandelana zihlala ngasikhathi sinye futhi zibambe iqhaza empilweni yekoloni.
Ukulandelana kokuziphendukela kwemvelo amazwe aphansi kwezenhlalo, okuholela ekwakhekeni kwezinga lokuhleleka kwenhlangano yezinambuzane, kumelelwa izintuthwane, izidakwa, iminyovu yokuzijabulisa kanye namaqembu ambalwa ezinyosi zenhlalo. Njengoba zonke izintuthwane eziphilayo nama-termite kuvuliwe izinga le-eusocial inhlangano yezenhlalo, ukuthuthukiswa okungaguquki kobudlelwano obungaphansi komhlaba kwafundwa kuma-wasps nezinye izinyosi. Kulomugqa wentuthuko, kukhona ukwanda kwezibopho phakathi kwabazali nenzalo. Ngendlela yokuphila eyedwa nasesigabeni esiphansi senhlalo, owesimame unakekela ukubekwa kwakhe phansi isikhashana, kepha ukuqunjelwa izingane kungalindi. Use Lesisemkhatsini wokuqala subsocial Esiteji, insikazi ihlala nabantu abasha ize ikhule. At the Lesisemkhatsini isigaba sesibili senhlalo ephansi abantu abadala abasha basiza abazali ekukhuliseni izingane ezintsha. Ukubambisana kuyabonakala phakathi kukamama nezingane, kepha hhayi phakathi kwendodakazi. Isigaba esilandelayo ukuvela kwamaqembu akhethekile abantu, abasebenzi, abasizi njalo ekukhulisweni kwesizukulwane esisha, lokhu izinga le-eusocial.
Ubungqabavu emikhakheni yezinambuzane ze-eusocial kuhamba ngezindlela ezimbili. Owokuqala ukukhulisa isibalo kanye nobungako bezinto ezikhethekile ezinxenyeni zabasebenzi. Umthombo wokwehlukanisa okwandayo kwezinqaba ezisebenzayo izinguquko eziqhubekayo zokuziphatha ezihlukile kubantu abahlukile, ngenxa yokuthi kwakheka izinhlobo ezahlukahlukene ze-morphological. Isibonelo, izintuthwane ezisebenza kahle zakha ikhanda elikhulu nama-mandibles, okuwavumela ukuba angenele amasosha. Kodwa-ke, ngenxa yezinguquko eziningi zokuziphatha kokuziphatha kumuntu munye, zingaba ngaphezu kokhokho olulodwa kukho konke impilo. Ezintuthwaneni I-Myrmica scabrinodis izisebenzi zesizini lokuqala lomsebenzi wazo ngemuva kokuqagela ezingqondweni zibamba iqhaza othandweni lwentsha, ngesizini ezayo iba ngabakhi, futhi nangemva kwalokho - ngabadali. Lokhu kuguqulwa kokusebenza kusebenza kahle kakhulu kwizinyosi zezinyosi.
Indlela yesibili yokukhethekile ezinambuzaneni ze-eusocial ukuthuthukiswa kokuxhumana phakathi kwabantu koloni, ukuze umsebenzi wabantu abaningi abahlala kuwo uhlelwe. Ukuxhumana, njengoba kuphawuliwe ekuqaleni kwalesi sahluko, kuthuthukiswa kakhulu phakathi kwezinambuzane zomphakathi. Ezinambuzaneni ze-eusocial, ukuxhumana kwamakhemikhali kuthuthukiswa kabanzi, kufaka phakathi ukukhishwa kanye nokuqonda kwamakhemikhali. Ngezinga elincane, kodwa futhi nokukhulumisana kahle kwamathekisthi athuthukiswe kahle, okukhona ukudonsa, ukushaya kanye nezinye izimpawu. Ukushintshwa kwama-pheromones e-liquid okucindezela ukwahlukaniswa kwe-Caste kungenye yezinto eziningi ezimangazayo eziphawulwa kumakoloni ezinambuzane.
Izinhlobonhlobo zezimpawu ezakha uhlelo lokuxhumana lwezinambuzane zihambelana nokusabela okuhlukahlukene kokuziphatha: ukukhathazeka, ukuheha kanye nokwakheka kwamaqoqo, kusesha imithombo yokudla okusha noma izindawo zezidleke, ukuqomisana, i-trophallaxis (ukushintshaniswa phakathi kwabantu abanezifo zomlomo noma zomchazi), ukudluliswa kwezinhlayiya zokudla kwabanye abantu, iqembu Ukuxhumana okwenyusa noma okunciphise umsebenzi ngamunye, ukukhonjwa kanye nokuqashelwa kwabalingani esidlekeni namalungu e-caste yabo, ukuzimisela kwabalingisi, okuvezwe njengo ukuvimbela, noma ukukhuthaza ukwahluka kwabo.
6.25. I-Colony yembovane yakudala ye-Australia bulldog (I-Myrmecia gulosa), akha isidleke sakhe emhlabathini
A. Uterus (Indlovukazi). B. Owesilisa. B. Isisebenzi esinikeza izifunisi zemilaba. Mnu Cocoons ngepupae.
6.26. Isidleke seThemu Amitermes hasatus
A. Iseli elingaphezulu ngezinsimu zokuzala. B. Iseli eliphakathi nendlovukazi - umsunguli wesidleke, owesilisa eduze kwakhe nabaningi abasebenzayo. B. Iseli eliphansi elinamasosha nama-nymphs lapho amasosha evela khona.
6.27. Imigqa emibili yokuthuthuka kwenhlangano ye-eusocial ezinambuzaneni zomphakathi - parasocial and subsocial
Ukungavikeleki Kwezinambuzane - Okujwayelekile
Ukuvikelwa kwezinambuzane ezifweni zamagciwane kuqinisekiswa bobabili ngokuba khona kwesembozo esiqinile se-chitinous, esisebenza njengesithiyo kwi-pathogen, kanye nokuba khona kokungavikeleki kwe-humoral kanye ne-cellular. Ucwaningo lwakamuva lukhombisa ukuthi izinambuzane azigcini nokugoma okungaphakathi kuphela, kodwa futhi zithole inkumbulo yokugonywa neyokugonywa.
Insect Cell Immunity
Ukungavikeleki kwamaselula kwezinambuzane kuvikela izinambuzane ezifweni ezibangelwa amagciwane ngokusebenzisa i-phagocytosis, ukubethela nokwakheka kwe-melanin kanye nokuphuma kwayo, okuyizinto ezinobuthi kumaseli wegciwane. Zonke lezi zinqubo zenzeka ngenxa yomsebenzi wezinhlobo ezintathu zamaseli: ama-plasmocytes, ama-lamellocyte, nokuhlanganisa amaseli we-phenol oxidase (amaseli we-crystal). Kwisinambuzane esidala, ama-plasmocytes ama-larval kuphela akhona, ngoba lesi sinambuzane silahlekelwa izindlala ze-lymph ngesikhathi se-metamorphosis kanye nesinambuzane esidala, amaseli we-immunocompetent awasakhiqizwa. Ku-izibungu zezinambuzane, zonke izinhlobo zamaseli we-immunocompetent zimelwe, kepha iningi lalaba bantu ngamaseli e-plasma. Amangqamuzana e-Phenol oxidase synthesizing cell akha i-5% kuphela yabantu bonke be-hemocyte. Ama-lammelocyte avela kuma-hemolymph wezibungu zezinambuzane kuphela lapho etheleleke nge-parasite enkulu, ama-plasmacyte angakwazi ukubhekana nayo. I-Phagocytosis yenziwa lapho i-plasmatocyte ibonwa ngomunye umuntu noma iguqulwa enye. Isibonelo, ama-phosphatidylserine aqukethe ama-phospholipids asendaweni yeseli esesimweni se-apoptosis. Abakwa-Plasmatocytes bababona besebenzisa ama-receptors athile futhi baqhuba i-phagocytosis. Uma i-ejenti yangaphandle engena emzimbeni wesinambuzane inkulu kakhulu, khona-ke ama-lamellocyte avela kubantu be-hemocyte - amaseli abandakanyeka enqubekweni yokuncipha. Ngakho-ke iminyovu yesinambuzane ibeka amaqanda hemocele we-Drosophila izibungu, ezihlaselwa ama-lamellocyte. AmaLamellocyte axhumeka ngaphezulu kweqanda futhi enza futhi ukuxhumana phakathi kwawo, akha i-capsule yemilayidi eminingi ezungeze iqanda le-parasite futhi ayihlukanise nemvelo yangaphakathi yomgcini. Futhi, amaseli ahlanganisa i-phenol oxidase ngaleyo ndlela ayakwazi ukukhipha i-oxidation yama-phenols kuma-quinones, lapho, lapho enza i-polymerized, enza i-melanin enobuthi kuma-microorganisms. Ngakho-ke, njengakwezilwane ezincelisayo, enye yezinqubo ezibalulekile zokungatheleleki kwamaselula ezinambuzaneni yi-phagocytosis eyenziwa yi-plasmatocytes. Ngakolunye uhlangothi, ngokungafani nezilwane ezincelisayo, izinambuzane ziyakwazi ukufaka engcupheni engase ibe khona esitshungwini, ngemuva kwalokho esingasuswa ndawo futhi zihlale emzimbeni wesinambuzane.
Ukungavikeleki kokulimazeka kwezinambuzane
Lapho amangqamuzana ezinambuzane angenamikhawulo esebenza ngokuhlangana kwamaphethini wamangqamuzana we-microbe, ama-receptor ahambisanayo ayasebenza bese kusayinwa amakhasethi ayabangelwa, okuholela ekusebenzeni kokubhaliwe kwezinhlobo eziningi zezakhi zofuzo nokutholakala kwamaprotheni asebenza njengama-antimicrobial agents. Ezinambuzaneni, izindlela ezimbili zokudlulisela amasignali zifundwa kangcono. Le ndlela eyi-Toll ibangelwa ukusebenzisana kwama-receptors ngesikhunta nangegciwane elinamandla egram (ngokuqondile, i-peptidoglycan yabo) kanye ne-Imd indlela ebangelwa ukusebenzisana kwama-receptors nge-peptidoglycan yamagciwane angen gamu. Njengomphumela wokwethulwa kwalezi zindlela zombili, inani lama-kinases we-intracellular lisebenza futhi isiginali etholwe nge-pathogen idluliselwa ku-nucleus. Ukwenza kusebenze i-ikB factor factor ye-ikB esimweni sokudluliswa kwesiginali nge-Toll signaling cascade kuholela ekuhambeni kwe-IkB kuyi-nucleus nasekubhalweni kwezakhi zofuzo zokulwa nokwelashwa.
Imikhiqizo Yokuqanjwa Kwezinambuzane ye-Gciwane
Ekuphenduleni ukutheleleka kuDrosophila, ama-peptides amafushane antimicrobial ahlanganiswa umzimba wamafutha nama-hemocytes. Abanye babo basebenza ngamagciwane angenayo i-gram-negative njenge-diptericin, abanye ngama-bacteria a-gram-positive afana ne-defensin kanye namagciwane fungal afana nokutheleleka kwe-Drosomycin. Ezinambuzaneni, amakilasi ayi-8 ama-peptides antimicrobial asevele abonakala, mhlawumbe kakhulu. Ngaphezu kwalokho, ama-peptides antimicrobial ayingxenye eyodwa yokuphendula kwesinambuzane ekuhlaselweni yi-pathogen. E-Drosophila, kuye kwatholakala izinhlobo zofuzo ezingama-543 okubhalwe ngazo okubhaliwe lapho kuphendulwa ukutheleleka. Imikhiqizo yokukhiqiza yalezi zakhi zofuzo yaziwa ngokuthi ama-pimides antimicrobial, cishe ama-25 ama-peptides angaziwa, amaprotheni abandakanyeka ekuqasheni kwamaphethini wamangqamuzana ebusweni be-pathogen kanye ne-phagocytosis, kanye namaprotheni abandakanyeka ekwenziweni kwezinhlobo ze-oksijini ezisebenzayo.
Amaprotheni e-DSCAM futhi athola impendulo yezinambuzane zokuzivikela
Ukwamukela ngokunembile noma yikuphi ukutheleleka okuye kwangena emzimbeni, kufaka phakathi okungakaze kuhlangane nakho ngaphambili, kufanele ube namaprotheni amaningi ahlukahlukene abopha ngokuthile ezintweni zangaphandle. AmaVertebrates axazulula inkinga yokubona omunye umuntu ongakabi nawo ukubhekana nokukhiqizwa kwamakhulu ezinkulungwane zama-antibody ahlukahlukene. Kuze kube muva nje, bekukholelwa ukuthi izinambuzane azinayo i-analogue yama-antibodies nokuthi kuphela ukuphendula kokuzivikela komzimba okungenzeka ezinambuzaneni. Kodwa-ke, ucwaningo lwakamuva luveze ukuthi mhlawumbe imikhiqizo ye-DSCAM gene ingahle ibambe iqhaza ekwakhiweni kwempendulo yokuzivikela yomzimba ezinambuzaneni. Uhlobo lwe-DSCAM lungokwamandla ama-immunoglobulins futhi ezinambuzini zibhekele ukulawulwa kokukhula kwe-axon. I-DSCAM iqukethe izivivinyo ezingama-21, kanti ezi-4, 6, 10 exons zimelwe ngabangu-14, 30, 38 amakhophi ngokulandelana. Njengomphumela wokuphinyiswa okunye, amaprotheni e-receptor ahlukene angama-15,960 angahlanganiswa. Ukuhlolwa komiyane womalaleveva kukhombisa ukuthi ukuvimba okwenziwe ngohlobo lwe-DSCAM kuholela ekwehleni kwekhono lomiyane lokumelana nezifo, futhi amagciwane aqala ukwanda ku-hemolymph yayo. Ngaphezu kwalokho, ukuhlukahluka kwe-splice kwe-DSCAM kunobuhlobo obukhulayo bobuso be-pathogen ekuphenduleni kokuhlaselwa okwenziwe ngabo. Ngakho-ke, ukuhlukahluka kwe-DSCAM kuphakamisa ukuthi badlala indima efanayo ezinambuzaneni njengama-antibodies kuma-vertebrates.
Lapho likhulu inani labantu ku-colony yezinyosi zenhlalo nezintuthwane, kuba buthaka ukungavikeleki kwabo.
Izazi zezinto eziphilayo zaseNyuvesi yaseNorth Carolina (e-USA) zithole ukuthi izinambuzane zomphakathi ezihlala emakoloni amaningi zinokuphendula okungatheni komzimba kokuvuselelwa kwamanye amazwe kunezihlobo zazo zakudala zenhlalo (ezihlala ngamaqembu amancane). Ososayensi bakholelwa ukuthi izinambuzane zenhlalo kungenzeka ukuthi zinezindlela ezithile ezingacacile ezivimbela ukusakazeka kwezifo, noma ngabe zivikela ukungatheleleki komzimba. Umsebenzi ushicilelwe ephephabhukwini. Izincwadi Zebhayoloji.
Sekukonke, kwahlolisiswa ukusabela kwezinhlobo eziyi-11 zezinambuzane, njengezenhlalo yezinyosi zezinyosi (Apis mellifera), amasimu (Zootermopsis nevadensis), izintuthwane ze-Woodworm (ICamponotus castaneus), futhi okungeyona eyokuhlalisana - ukuhlala izinyosi zodwa, iminyovu kanye namaphela.
Ukuhlola umsebenzi wokungavikeleki kwabo, ababhali bomsebenzi bazame ukuvusa impendulo yokuzivikela ezifweni ezihlolwayo. Ukuze benze lokhu, basebenzise i-probe eyethulwa emzimbeni wama-arthropods angenayo i-polypysaccharides micron imicu yenayiloni enobude bamamilimitha amathathu.Emvelweni, i-lipopolysaccharides iyisakhi esiyinhloko sodonga lweseli lwamagciwane angenawo i-gram-negative, ngakho-ke ukungatheleleki kwezifo eziningi kuthatha ukwakheka okunjalo njengama-ejenti angenasifo futhi aqala ukuwahlasela. Ngemuva kwenkathi yama-incubation yamahora amane, iphenyi elinentambo yenayiloni lahoxiswa emuva futhi umbala wayo wathathwa izithombe.
Iqiniso liwukuthi amasosha omzimba ezinambuzane asebenza ngokuphumelelayo ukulwa nezifo: lizungeza umzimba ongaphandle 'ngodonga' lwama-hemocytes (izifanekiselo ezikude zamangqamuzana egazi abantu nama-lymphocyte). Ama-hemocytes amaningi ayesentanjeni, i-melanin eyengeziwe yayikuyo futhi mnyama mnyama ngombala wawo ngemuva kokuhlolwa.
Kwavela ukuthi phakathi kwezinambuzane zomphakathi nezihlala zodwa akunemehluko omkhulu ngezimpendulo zokuzivikela komzimba. Kepha ngaphakathi kweqembu lezilokazane zenhlalo impendulo yokuzivikela komzimba yayingaphansi kunasemikhakheni emikhulu ababehlala kuyo. Ngakho-ke, ukungavikeleki kokugomela kwabonwa ezinyosini zezinyosi ngemikoko yazo emikhulu, nasezinyosi zasemhlabeni (IHalictus ligatus), amakoloni ayo anezakhamizi ezimbalwa, impendulo yokuzivikela komzimba yayinamandla amakhulu.
Okwamanje, kuba nengxoxo emphakathini wesayensi mayelana nokuthi izinambuzane zomphakathi zihlangabezana kanjani ngqo nosongo lwezifo eziwumshayabhuqe. Imvamisa, izidalwa eziphilayo zigwema amaqembu amakhulu ezinhlobo zazo, ngoba ezindaweni ezinjalo kunamathuba aphezulu okuthola isifo esithathelwanayo. Izinqubo ezifanayo zingavezwa kalula ngesibonelo sabantu ababengasazi isifo esikhulu se-Neolithic, kepha ngubani nge-Iron Age ovame ukulahlekelwa yinqwaba yabantu bendawo ethile evela kubo.
Kuze kube manje, amaphuzu amabili okubukwa abekwe phambili ukuthi izinyosi, izintuthwane, kanye nezinye izinambuzane ezifanayo kugwema kanjani ukulahleka okukhulu kwezinhlobo zabantu ezifweni. Ngokusho kokuqala, kumane kunezifo eziqinile zokugomela, ezibonakala ziphikisana kakhulu nesizinda esijwayelekile sezinambuzane. I-hypothesis yesibili yathi ukungavikeleki kwabo kuyinto evamile, kepha izinambuzane zenhlalo zithuthukise izindlela ezinciphisa ubungozi bokutheleleka noma ukudluliselwa, ngokwesibonelo, ukuhlanzeka okuthuthukile. Kumele kwaziwe ukuthi inyosi efanayo yezinyosi ihlala izihlanza, futhi uma inephunga, ingahle ingavunyelwa ukungena esidlekeni yizihlobo zayo.