Pakati pa tizilombo, mitundu yovuta kwambiri yamakhalidwe imawonedwa mu tizilombo ta gulu. Bungwe la madera awo limakhazikitsidwa chifukwa cha ubale ndi mgwirizano womwe umakhalapo pakati pa mamembala ake omwe amakhala mdera limodzi ndikugawana chisa cha mtundu wina. Zilombo zenizeni zachikhalidwe, zomwe zimatchedwa eusocial tizilombo, ndi am'magulu awiri. Awa onse ndi malovu (Isoptera) ndi nthumwi zapadera za Hymenoptera. nyerere zonse ndi zina zoyipitsidwa bwino kwambiri ma njuchi ndi njuchi.
Khalidwe la Eusocial limadziwika ndi zinthu zazikulu zitatu.
- m'malire amtunduwu, anthu amaphatikiza kusamalira ana,
- ntchito pagulu limagawidwa m'magulu apadera aanthu,
- magawo amoyo a anthu osachepera awiri mibadwo imachulukana, kotero kuti mbadwo wachichepere umakhala gawo limodzi la moyo wawo ndi mbadwo.
Pazigawo za gulu la pre-social (presocial) gulu la tizilombo, chimodzi kapena ziwiri zokha mwa izi zitatu ndizikhalidwe.
Zilombo zamtundu wa Eusocial zimadziwikanso ndi kuchuluka kwambiri kwa kubereka. Chiwerengero chaanthu omwe ali mgululi chimachokera ochepera 100 mpaka mamiliyoni ambiri amitundu mitundu ndi magulu a tizilombo (mkuyu. 6.25, 6.26). Danga limodzi la mitundu ya nyerere zaku Africa limatha kukhala ndi anthu pafupifupi 22 miliyoni. Ena mwa nyerere zokonzedwa bwino kwambiri Myrmica rubra, m'dulamo mumakhala chiberekero chimodzi ("mfumukazi"), chomwe chimakhala zaka zingapo, chikuikira mazira ndi kubereka ana kumudzi wonsewo, ndipo akazi onse osabereka amapereka chakudya. Anthu osabereka amagwira ntchito zambiri munthaka.
Nthawi zambiri amaimiridwa ndi mitundu yosiyanasiyana ya morphological, kapena castes. Anthu ochepa kwambiri - ogwira ntchito, zokulirapo - asirikali kapena antchito akulu. Ogwira ntchito amatenga nawo mbali popeza chakudya, chisamaliro, kusamalira ana. Asitikali amateteza chisa komanso malo osungira zakudya zamadzimadzi. Anthu osabereka amakhala ndi moyo waufupi, ndipo chiberekero chimayenera kuyikira mazira mosalekeza kuti zitsimikizire kuti njuchiyo ndi yayikulu panthawi yakukhwima. Kuphatikiza pa magulu amtunduwu, pali gulu lina lomwe lili amuna. Sagwira ntchito iliyonse, amakhala ndi zochitika zina zokomera ena (mwachitsanzo, kusamalira anthu ena) ndipo akuyembekezera kuthawa kwawo kuti akwaniritse akazi achikazi. Zazikazi zazikazi zimachokera ku mbadwa za wopanga wamkazi. Nyengo ikamatha, aliyense wamkazi amakhazikitsa gulu lake latsopano, ndikumanga chisa ndi kusamalira ana. Mbewu ikadzakhwima, anthu osabereka amasamalira ana ndi ntchito zina.
Zomwe zidasinthika kukhala gawo la eusocial gulu la tizilombo zinali mizere iwiri yolimbikitsa ubale - parasocial ndi ochezeka (Mkuyu. 6.27). Ndi njira yokhayokha (yopanda chikhalidwe) yokhala ndi ana, palibe chodera nkhawa ana, palibe njira zapadera zoberekera komanso kuphatikizana kwa mibadwo yotsatizana. Ndi gulu la parasocial lodziwika bwino la njuchi za banja la a Halictidae, anthu akuluakulu am'badwo umodzi, pamlingo wina kapena wina, amathandizana. Gawo lotsika kwambiri la bungwe lamtunduwu limatchedwa wamba. M'madera omwe ali ndi gawo ili, akuluakulu amakumana kuti amange chisa, koma ana amaleredwa payokha.
Pa gawo lachitukuko pali mitundu ingapo ya njuchi yochokera ku banja la a Halictidae. Pafupifupi azimayi 50 amtundu womwewo amakhala mchisa pansi. Njuchi iliyonse imakumba m'mphepete mwake kapena maselo am'modzi, pomwe imagwiritsa ntchito dzira limodzi, imayika chakudya ndikumatula. Mulingo wotsatira wa mzere wa parasocial - kopitilira muyeso - Kusamalira ana ndi kwa anthu, koma mkazi aliyense amaikira mazira panthawi yake. Lotsatira, hemisocial, wogwira antchito amawonekera, wopangidwa ndi anthu omwe sanali kubereka khola. Gawo lotsatira la mabungwe kale zosavutaomwe amakwaniritsidwa ngati chiyembekezo cham'badwo wam'badwo wa hemisocial chikhala chachikulu kwambiri mwakuti mibadwo iwiri kapena kupitilira apo imakhala nthawi imodzi ndikuchita limodzi moyo wa gulu.
Kusinthika kwotsatira mayiko ocheperako, zomwe zikuwatsogolera pakupanga gulu la tizilombo tosiyanasiyana, timayimiridwa ndi nyerere, chiswe, mavu ochezera komanso magulu angapo a njuchi zothandizirana. Popeza nyerere zonse zokhala ndi chonde zili mulingo wokongola bungwe lazachikhalidwe, kukula kosalekeza kwa ubale wapansi kunaphunziridwa pa mavu ndi njuchi zina. Mu mzere wa kakulidwe kameneka, pali kuwonjezeka kwa ubale pakati pa makolo ndi ana. Ndi moyo wapayekha komanso pamalo oyamba okhalamo, mzimayi amasamalira kugona komwe amakhala pansi kwakanthawi, koma samadikirira kuwonongedwa kwa achinyamata. Pa wapakatikati woyamba wocheperako Pa siteji, wamkazi amakhalabe ndi achinyamata mpaka atakula. Pakatikati gawo lachiwiri lokondwerera Achinyamata akuluakulu amathandizira makolo polera mwana watsopano. Mgwirizano umawonedwa pakati pa mayi ndi ana, koma osati pakati pa mwana wamkazi. Gawo lotsatira ndikutuluka kwa magulu apadera aanthu, ogwira ntchito, omwe amakhala othandizira pakukweza kwa m'badwo watsopano, izi ndi mulingo wokongola.
Kukhazikika mu magulu a tizilombo ta eusocial kumachitika m'njira ziwiri. Loyamba ndikuwonjezera chiwerengero ndi madigiri a akatswiri mu antchito. Zomwe zimawonjezera kusiyanasiyana kwa mabwalo omwe amagwira ntchito ndizosintha mosasintha m'makhalidwe omwe ali osiyana m'magulu osiyanasiyana, chifukwa chake mitundu yosiyanasiyana ya morphological imapangidwa. Mwachitsanzo, nyerere zantchito zabwino zimapanga mutu waukulu ndi mandibles, zomwe zimawalola kulowa gulu la asitikali. Komabe, chifukwa cha kusintha kwachilengedwe kwa munthu m'modzi, kumatha kukhala kwamunthu wopitilira umodzi m'moyo wonse. Pa nyerere Myrmica scabrinodis ogwira ntchito mu nyengo yoyamba ya zochitika zawo ataganiza molting atatenga nawo mbali pachibwenzi cha achinyamata, mu nyengo yotsatira amakhala opanga, ndipo ngakhale pambuyo pake - forageers. Kusintha kwa ntchito kumeneku kumapangidwa bwino kwambiri mu njuchi za uchi.
Njira yachiwiri yosinthira tizilombo ta eusocial ndikutukuka kwa kulumikizana pakati paanthu omwe ali mgululi, kuti zochita za anthu ambiri omwe akukhalamo zigwirizane. Kulankhulana, monga tanena kale m'mutu uno, zimapangidwa kwambiri pakati pa tizilombo tokhala pagulu. Mu tizilombo ta eusocial, kulumikizana kwamankhwala kumapangidwa kwambiri, kuphatikiza kumasulidwa ndi kuzindikira kwa mankhwala. Pocheperako pang'ono, komanso kulumikizana bwino kwamayendedwe, komwe kumangogunda, kugogoda ndi zizindikilo zina. Kusinthana kwa ma pheromoni amadzimadzi omwe amapewetsa kusiyanasiyana kwa ma caste ndi chimodzi mwazinthu zambiri zodabwitsa zomwe zimawonedwa m'mazilombo.
Zosiyanasiyana zomwe zimapanga njira yolumikizirana ya tizilombo tofanana ndi machitidwe osiyanasiyana: nkhawa, kukopa ndikupanga masango, kusaka magwero atsopano azakudya kapena malo a zisa, chibwenzi, trophallaxis (kusinthanitsa pakati pa anthu omwe ali ndi milomo yamkamwa kapena anal fluid), kusamutsa chakudya kwa anthu ena, gulu kulumikizana komwe kumalimbitsa kapena kufooketsa zochita za aliyense payekha, kuzindikira ndi kuzindikira za abwenzi mu chisa ndi mamembala awo, kutsimikiza kwa ochita masewera, omwe akuwonetsedwa ngati poletsa, kapena kukondoweza kwa kusiyana kwawo.
6.25. Colony wa nyerere wakale waku Australia bulldog (Myrmecia gulosa,, akumanga chisa chake munthaka
A. Uterus (Mfumukazi). B. Amuna. B. Wogwira ntchito yopereka mphutsi. Mr. Cocoons okhala ndi pupae.
6.26. Chisa chogona Amitermes hasatus
A. Upper cell yokhala ndi nymphs. B. Selo yapakati ndi mfumukazi - woyambitsa chisa, wamwamuna pafupi ndi iye ndi anthu ambiri ogwira ntchito. B. Selo yotsika ndi asitikali ndi ma nyezi omwe asitikali amasintha.
6.27. Mizere iwiri ya chitukuko cha eusocial gulu pagulu tizilombo - parasocial ndi subsocial
Kuteteza Tizilombo - General
Kuteteza tizilombo ku matenda oyambitsidwa ndi kachilombo ka kachilomboka kamafotokozedwa ndi kupezeka kwa chivundikiro cholimba cha chitinous, chomwe chimalepheretsa tizilombo toyambitsa matenda, komanso kukhalapo kwa chitetezo chazovuta komanso ma cell. Kafukufuku waposachedwa akuwonetsa kuti tizilombo samangokhala ndi chitetezo chokwanira chokha, komanso chitetezo cha m'thupi komanso chitetezo cha m'thupi.
Tizilombo Toyambitsa Matenda
Kukhazikika kwa chitetezo cha ma cell kumateteza tizirombo toyambitsa matenda kuchokera ku ma phagocytosis, kufalikira komanso kapangidwe kake ka melanin ndi zotumphukira zake, zomwe ndi poizoni m'maselo a bakiteriya. Njira zonsezi zimachitika chifukwa cha ntchito ya mitundu itatu ya ma cell: ma plasmocytes, lamellocyte, ndikupanga maselo a phenol oxidase (maselo a kristalo). Mu tizilombo toyambitsa matenda akuluakulu, tizilombo tating'onoting'ono tokha timene timapezeka, chifukwa tizilombo timene timataya timadzi tomwe timatulutsa timadzi tambiri pa metamorphosis komanso tizilombo toyambitsa matenda akuluakulu, maselo osakwanira sapangidwanso. Mu mphutsi zamtundu, mitundu yonse ya maselo a immunocompetent amaimiridwa, komabe, ambiri mwa anthuwa ndi plasmatocytes. Maselo ophatikizira a Phenol oxidase amapanga 5% yokha ya hemocyte yonse. Lammelocyte amapezeka mu hemolymph wa tizilombo tokhala ndi tizilombo tating'onoting'ono pokhapokha titakhala ndi majeremusi akuluakulu, omwe ma plasmacyte sangathe kupirira. Phagocytosis imachitika pamene plasmatocyte imadziwika ndi wina kapena wasintha. Mwachitsanzo, ma phosphatidylserine okhala ndi ma phospholipids ali pachiwonetsero cha cell. Plasmatocytes amawazindikira akugwiritsa ntchito ma receptor enieni ndikuchita phagocytosis. Ngati wogulitsa wakunja akulowa m'thupi la tizilombo ndiochuluka kwambiri, ndiye kuti ma lamellocyte amawonekera mu hemocyte kuchuluka - maselo omwe akukhudzidwa ndi njira yowonjezera. Chifukwa chake mavu a parasitic amayikira mazira mu hemocele ya Drosophila mphutsi, zomwe zimatsutsidwa ndi lamellocyte. Ma Lamellocyte amalumikizidwa pamwamba pa dzira ndipo amapanganso kulumikizana pakati pawo, ndikupanga kapikisano kazinthu zingapo zomwe zimazungulira dzira la kachiromboka ndikudzilekanitsa ndi chilengedwe chamkatikati. Nawonso maselo omwe amapanga phenol oxidase amatha kuyambitsa makutidwe amtunduwu kuti asungidwe, zomwe, zikapangidwa, zimapangika melanin poizoni tizilombo tosiyanasiyana. Chifukwa chake, monga momwe ziliri kwa zolengedwa zazikazi, imodzi mwazinthu zazikulu kwambiri za chitetezo chokwanira mu tizilombo ndi chiwgocytosis zimachitika ndi plasmatocytes. Komabe, mosiyana ndi zinyama, tizilombo timatha kupaka vuto lomwe lingakhale pachiwopsezo, chomwe pambuyo pake sichichotsedwa kwina konse ndipo chimakhalabe m'thupi la kachilombo.
Kuthetsa chitetezo chokwanira kwa tizilombo
Maselo atizilombo tating'onoting'ono tomwe timalumikizana ndi ma cell maselo okhala ndi michere, ma receptor ofanana amawayambitsa ndikuwonetsa ma cascade amayambitsidwa, zomwe zimayambitsa kutsegula kwa mitundu yambiri ya antimicrobial komanso kuphatikiza mapuloteni omwe amagwira ntchito ngati antimicrobial agents. Mu tizilombo, njira ziwiri zotumizira bwino zimaphunziridwa bwino. Iyi ndiye njira ya Toll yomwe imayambitsidwa ndi kulumikizana kwa ma receptors okhala ndi bowa ndi mabakiteriya abwino (moyenera, peptidoglycan) ndi njira ya Imd yomwe imayambitsidwa ndi kulumikizana kwa receptors ndi peptidoglycan ya mabakiteriya opanda gramu. Zotsatira zakukhazikitsidwa kwa njira zonse ziwirizi, zigawo zingapo zama intracellular zimayambitsidwa ndipo chizindikiro cholandila za pathogen chimapititsidwa ku khutu. Kukhazikitsidwa kwa cholembera cholembedwa cha ikB pankhani ya kufalikira kwa chizindikiritso kudzera pa Toll signaling cascade kumabweretsa kuyenda kwa IkB mu nyukiliya ndikulemba mitundu yotsutsana ndi antimicrobial.
Tizilombo Togulitsa Tizilombo toyambitsa matenda a Gene
Poyankha matenda ku Drosophila, ma peptides amtundu waufupi amapangidwa ndi thupi lamafuta ndi hemocytes. Ena mwa iwo amakhala ndi mabakiteriya osagwiritsa ntchito gramu ngati diptericin, ena pamabakiteriya okhala ndi gramu monga defensin ndi mabakiteriya oyambira ngati drosomycin. Mu tizilombo, magulu asanu ndi atatu a ma peptides opangira antimicrobial adadziwika kale, mwina ochulukirapo. Kuphatikiza apo, ma peptides antimicrobial ndi gawo limodzi lokha pazomwe tizilombo timayambitsa pakuwukira kwa pathogen. Ku Drosophila, majini a 543 adadziwika omwe amalemba omwe adawongolera poyankha matenda. Zinthu zomwe zimapangidwa pamtunduwu zimadziwika kuti zimayambitsa matenda opha tizilombo, pafupifupi ma 25 peptides osadziwika, mapuloteni omwe amathandizira kuzindikira mawonekedwe a maselo padziko lapansi la pathogen komanso phagocytosis, komanso mapuloteni omwe amathandizira kupanga mitundu ya oxygen yomwe imathandizira.
DSCAM mapuloteni komanso kupeza tizirombo toyambitsa matenda
Kuti muzindikire bwino matenda aliwonse omwe alowa mthupi, kuphatikiza omwe sanakumanepo nawo, muyenera kukhala ndi mapuloteni osiyanasiyana omwe amasankha mosiyana ndi zinthu zakunja. Ma Vertebrates amathetsa vuto lakudziwitsa munthu wina yemwe sanachite nawo ntchito yopanga mazana a masauzande ambiri a antibody. Mpaka posachedwa, anthu amakhulupirira kuti tizilombo toyambitsa matenda sitingathe kudziwa momwe tingagwiritsire ntchito tizilombo tating'onoting'ono. Komabe, kafukufuku waposachedwa akuwonetsa kuti mwina mankhwala a geni a DSCAM atha kutenga nawo mbali popanga yankho la chitetezo cha m'thupi la tizilombo. Mtundu wa DSCAM ndi wa superunamulopulini ndipo mwa tizilombo timayambitsa chiwongolero cha axon. DSCAM ili ndi ma exons 21, ndipo 4, 6, 10 exons oyimiridwa ndi 14, 30, 38 makope, motsatana. Chifukwa cha kugawanika kwina, mapuloteni osiyana-siyana a 15,960 amatha kupanga. Kafukufuku wokhudza udzudzu wa malungo adawonetsa kuti kubisa kwa jini la DSCAM kumabweretsa kuchepa kwa mphamvu ya udzudzu kukana matenda, ndipo mabakiteriya amayamba kuchulukana mu hemolymph yake. Kuphatikiza apo, ma splice osiyanasiyana a DSCAM ali ndi mgwirizano wowonjezereka wa pathogen poyankha momwe adapangidwira. Chifukwa chake, kusiyanasiyana kwa DSCAM kukuwonetsa kuti amatenganso mbali zofanana ndi tizilombo tofanana ndi ma antibodies a vertebrates.
Kuchulukana kwa anthu omwe ali ndi njuchi ndi nyerere, kumachepera chitetezo chawo.
Akatswiri azachilengedwe ochokera ku Yunivesite ya North Carolina (USA) apeza kuti tizilombo tomwe timakhala m'madera ambiri sachitapo kanthu poyang'anira zikhalidwe zakunja kuposa abale awo oyamba (okhala m'magulu ang'onoang'ono). Asayansi akukhulupirira kuti tizilombo tomwe timakhala pagulu mwina tili ndi njira zina zowoneka bwino zomwe zimalepheretsa kufalikira, ngakhale atachepetsa chitetezo chokwanira. Ntchitoyi idasindikizidwa mu mtolankhani. Makalata a Biology.
Pazonse, mitundu ya mitundu 11 ya tizilombo, monga njuchi zanjuchi, idaphunziridwa (Apis mellifera), chiswe (Zootermopsis nevadensis), nyerere zamtchire (Camponotus castaneus, komanso osakhala ochezeka - amakhala okha njuchi, mavu ndi maphemwe.
Kuti awone ntchito ya chitetezo chawo, olemba ntchitoyi adayesetsa kuyambitsa chitetezo cha mthupi pazoyeserera. Kuti achite izi, pogwiritsa ntchito kafukufuku, adayambitsa ma microfilaments atatu atali a mamililita ophatikizidwa ndi lipopolysaccharides kulowa mu thupi la mankhwala osagwiritsidwa ntchito ndi lipopolysaccharides.Mwachilengedwe, lipopolysaccharides ndiye gawo lalikulu la khungu la mabakiteriya osagwiritsa ntchito, motero kusatetezeka kwa zinthu zambiri kumachitika ngati mankhwala opatsirana ndikuyamba kuwayambitsa. Pambuyo pakupanga kwa maola anayi, chifanizo chokhala ndi ulusi wa nayiloni chinachotsedwa ndipo mtundu wake udali kujambulidwa.
Chowonadi ndi chakuti chitetezo cha mthupi cha tizilombo chimagwiritsa ntchito kulowetsa matenda pofuna kuthana ndi matenda: imazungulira thupi lakunja ndi "khoma" la hemocytes (fanizo lakutali la maselo amagazi a anthu ndi ma lymphocyte). Ma hemocytes ochulukirapo omwe anali ndi ulusi, momwemonso ma melanin amapezeka ndipo amdima ake anali amdima utatha kuyeserera.
Zidapezeka kuti pakati paanthu ndi tizilombo tokhala patokha palibe kusiyana kwakukulu pakayankho ka chitetezo cha mthupi. Koma mkati mwa gulu la tizilombo tating'onoting'ono, mayankho a chitetezo mthupi anali ochepa mphamvu poyerekeza ndi magulu akuluakulu omwe amakhala. Chifukwa chake, kuponderezedwa chitetezo chokwanira kumawonedwa ndi njuchi zazing'ono ndi ming'oma yawo yayikulu, komanso njuchi zapadziko lapansi (Halictus ligatus), omwe madera ake ali ndi anthu ochepa, chitetezo cha mthupi chidalimba kwambiri.
Pakadali pano, pali zokambirana pagulu la asayansi zokhudzana ndi momwe ndudu zapagulu zimatha kuthana ndi vuto la miliri. Nthawi zambiri, zinthu zachilengedwe zimapewa masango akuluakulu amtundu wawo, chifukwa m'malo otere pali mwayi waukulu wodwala matenda opatsirana. Njira zofananazi zitha kufotokozedwa mosavuta ndi zitsanzo za anthu omwe sanadziwebe miliri yayikulu mu Neolithic, koma ndi Iron Age nthawi zambiri amataya kuchuluka kwa anthu amderali kuchokera kwa iwo.
Mpaka pano, mbali ziwiri za malingaliro zakambidwa momwe ndendende, nyerere, ndi tizilombo tina tomwe timapewera kuchepa kwakukulu kwa chiwerengero kuchokera kumatenda. Malinga ndi oyambayo, amangokhala ndi chitetezo chathupi champhamvu kwambiri, chomwe chimadziwika kwambiri kuyambira kale ngati tizilombo. Hypothesis yachiwiri idati kusatetezeka kwawo ndikwabwinobwino, koma tizilombo tomwe timakhala pagulu tapanga njira zomwe zimachepetsa chiopsezo chotenga kachilomboka kapena kufalitsa, mwachitsanzo, ukhondo wopitilira. Dziwani kuti njuchi yomweyo imadziyeretsa, ndipo ngati ikununkhira, singalole kulowa mng'oma ndi abale ake.