Chim

Thi im ha thch: 1500 Ma

Cui k Jura - nay

Bch khoa ton th m Wikipedia

Chim (danh php khoa hc: Aves ) l tp hp cc loi ng vt c xng sng, mu nng, i ng bng hai chn v trng. Trong lp Chim, c khong 10.000 loi cn tn ti, gip chng tr thnh lp a dng nht trong cc loi ng vt bn chi. Lp chim c tr cc h sinh thi khp ton cu, t vng Bc Cc cho ti chu Nam Cc. Cc loi chim c kch thc dao ng khc nhau, t nh c 5 cm (nh Mellisuga helenae - mt loichim rui) cho ti ln c 2,7 m (nh iu). Cc bng chng ha thch cho thy, chim c tin ha t cc loi khng long chn th(Theropoda) trong sut k Jura, vo khong 150-200 triu nm v trc, vi i din u tin c bit n, xut hin t cui k Jura lArchaeopteryx (vo khong 155150 triu nm trc). Hu ht cc nh c sinh vt hc u coi chim l nhnh duy nht ca khng long cn sng st qua s kin tuyt chng CretaceousTertiary vo xp x 65,5 triu nm trc. Cc loi chim hin i mang cc c im tiu biu nh: c lng v, c m v khng rng, trng c v cng, ch s trao i cht cao, tim c bn ngn, cng vi mt b xng nh nhng chc. Tt c cc loi chim u c chi trc bin i thnh cnh v hu ht c th bay, tr nhng ngoi l nh cc loi thuc b Chim cnh ct, b iu v mt s a dng nhng loi chim c hu sng trn o. Chim cng c h tiu ha vh hp c nht m p ng cao cho hot ng bay. Vi loi chim, c bit l h Qu v vt, nm trong nhng loi thng minh nht ca gii ng vt; mt s c quan st ang ch to v s dng cng c, nhiu loi sng thnh by li c th truyn t nhng kinh nghim hiu bit cho th h sau. Nhiu loi chim hng nm thng di tr n nhng ni rt xa, cng rt nhiu loi li thc hin nhng chuyn bay ngn hn v bt thng. Chim l ng vt sng by n, chng giao tip vi nhau thng qua ting ku v ting ht, tham gia vo nhng hot ng by n nh hp tc trong vic sinh sn, sn mi, di chuyn v tn cng chng li k th. Phn ln chim l nhng loi n giao x hi[1], thng vo ma giao phi trong mt thi gian nht nh. Trng chim thng c trong t v p bi chim b m. Hu ht chim non sau khi n u c thm mt thi gian c chim b m chm sc. Nhiu loi chim c tm quan trng i vi con ngi, a phn c s dng lm thc n thng qua vic sn bn hay chn nui. Mt vi loi, nhphn b S hay b Vt, c bit n vi vai tr vt nui lm cnh. Hnh tng chim xut hin trong tt c cc mt ca vn ha con ngi, t tn gio, thn thoi n thi ca v m nhc ph thng. Khong 120-130 loi chim b tuyt chng do hnh ng con ngi trong th k 17, cng vi hng trm loi khc sau . Hin nay, c khong 1.200 loi ang trong tnh trng e da tuyt chng bi cc hot ng t loi ngi, cho d vn ang c nhng n lc bo v chng.

Tin ha v phn loi

Archaeopteryx, i din sm nht ca chim c bit n.

Chim c phn loi ln u tin bi Francis Willughby v John Ray, a ra trong tp sch Ornithologiae nm 1676[2]. Carolus Linnaeus sau sa i cng trnh ny vo nm 1758 v li h thng phn loi vn cn c s dng cho n ngy nay[3]. Trong h thng phn loi Linnaeus, cc loi chim c phn vo lp Aves. Vic phn loi da trn ngun gc pht sinh xp Aves vo trong nhnh khng long Theropoda (Khng long chn th)[4]. Aves cng vi nhm ch em Crocodilia (B C su) l nhng thnh vin duy nht cn sng st ca nhnh b st Archosaur. Xt theo pht sinh chng loi, Aves c xc nh l tt c cc hu du ca t tin chung gn nht ca cc loi chim hin i v Archaeopteryx lithographica[5]. Theo nh ngha ny, Archaeopteryx, xut hin t giai on Tithonia ca Hu Jura (150-145 triu nm trc), l chi chim sm nht c bit n. Nhng lun im khc, bao gm Jacques Gauthier v nhng ngi ng h h thng PhyloCode, xc nh Aves ch bao gm cc nhm chim hin i, ch khng gm hu ht cc nhm chim ch c bit n qua ha thch, v thay vo xp chng vo nhm ln Avialae[6], mt phn trnh c s khng r rng v v tr ca Archaeopteryx trong gii ng vt m theo quan im truyn thng thng coi chng l nhng khng long chn th. Tt c cc loi chim hin i u nm trong phn lp Neornithes, chia thnh hai nhm: Paleognathae, bao gm hu ht cc loi khng bit bay nh iu, v Neognathae a dng hn, cha tt c nhng loi cn li[4]. Hai nhm ny thng c xp cp bc siu b[7], d Livezey & Zusi t chng cp "cohort"[4]. Ty vo quan im phn loi khc nhau, s lng loi chim cn tn ti dao ng t 9.800[8] cho n 10.050 loi[9].

Khng long v ngun gc ca chim

Cc bng chng ha thch v phn tch sinh hc chuyn su chng minh vt qua bt c nghi ng c l no rng chim l nhng loi khng long chn th. C th hn, chng l thnh vin ca Maniraptora, cng vi cc nhm khc nh h Dromaeosauridae v h Oviraptoridae[10]. Khi cng nhiu loi khng long chn th c mi quan h gn gi vi chim c cc nh khoa hc pht hin, th nhng im khc bit r rng trc y gia chim v loi khng phi chim li cng b xa nha. Nhng pht hin gn y ti tnh Liu Ninh pha ng Bc Trung Quc, chng t rng c nhiu khng long chn th c nh c lng v, li cng gp phn thm cho s khng r rng ny[11]. Quan im ng thun trong gii c sinh vt hc hin i cho rng lp Chim (Aves), chnh l h hng gn gi nht ca cn b Deinonychosauria, m bao gm hai h Dromaeosauridae vTroodontidae. Chng cng nhau to thnh mt nhm gi l Paraves. chi c s Microraptor ca h Dromaeosauridae, cng c b lng v m c th c chng s dng bay ln. Hu ht nhng loi Deinonychosauria c s u rt nh b; iu ny gia tng kh nng rng t tin ca tt c cc loi Paraves c th tng sng trn cy, v/hoc c kh nng chao ln[12][13]. Archaeopteryx ca thi k Hu Jura c bit n nh l mt trong nhng ha thch chuyn tip u tin c tm thy, v iu ny cng gip ng h thm cho hc thuyt tin ha vo cuith k 19. Archaeopteryx c nhng c im ca ng vt b st nh: c rng, tay c mng vut, cng mt chic ui di v ging thn ln, tuy nhin n cng c trang b ring mt i cnh tinh vi vi nhng chic lng bay m ging ht nh nhng con chim hin i. N khng c cng nhn l t tin trc tip ca chim hin i, nhng vn l thnh vin c xa v nguyn thy nht ca nhm Aves hay Avialae, v gn nh chc chn l c mi quan h gn gi vi chim t tin thc s. Tuy vy, vn c gi thuyt cho rng Archaeopteryx thc s l khng long v khng c mi quan h gn gi no hn vi chim so vi cc nhm khng long khc[14], v rng Avimimus cn ging vi t tin cc loi chim hn c loi ny[15].

Cc thuyt v tranh lun khc

c rt nhiu tranh lun n ra v ngun gc cc loi chim. Khi u, ngi ta tranh lun v chim bt ngun t khng long hay t nhng Archosaur c xa hn. Trong pha ng h gi thuyt khng long, cng c nhng bt ng v Ornithischia hay khng long chn th mi gn hn vi nhng loi thy t[16]. D Ornithischia (khng long "hng chim") c cu trc hng tng ng vi nhng loi chim hin i, chim vn c coi l bt ngun t ging khng long Saurischia ("hng thn ln"), m tin ha cu trc hng ca chng mt cch c lp[17]. Trn thc t, bn

cnh hai nhm trn, cu trc hng ging chim cng c tin ha trong mt nhm khng long chn th ring bit, c bit n l Therizinosauridae. Hai nh khoa hc Larry Martin v Alan Feduccia tin rng chim khng phi l khng long, m tin ha t nhng loi Archosaur s khai nh Longisquama. Lun im chnh trong cng b ca h cho rng nhng tng ng gia chim v khng long Maniraptora thc t l do s ng quy, v c hai khng lin quan g ti nhau. Vo cui nhng nm 1990, bng chng v chim l Maniraptora tr nn khng th chi ci, nn Martin v Feduccia chp nhn phin bn sa i trong gi thuyt ca h bi ngh s dn dng khng long Gregory S. Paul; trong nhng Maniraptora l nhng loi chim khng bit bay th cp[18], tuy nhin trong phin bn , chim li tin ha trc tip t Longisquama. Theo nh vy th chim vn khng phi khng long, nhng cng khng phi l hu ht cc loi c bit n m hin ti c phn loi nh khng long chn th. Maniraptora, thay v th, li l nhng loi chim thuc nhm Archosaur v khng bit bay[19]. Gi thuyt ca Martin v Feduccia khng c tha nhn bi hu ht cc nh c sinh vt hc.[20]. Nhng c im m xem nh bng chng ca s khng bay c, theo xu th ch o ca nhng nh c sinh vt, c gii thch l "s thch nghi ban u" (pre-adation hay exaptation), rng Maniraptora tha hng nhng c im t t tin chung ca chng cng vi cc loi chim. Protoavis texensis c miu t nm 1991, coi l loi chim c hn c Archaeopteryx. Tuy nhin, nhng ngi ch trch cho rng ha thch pht hin c cht lng bo qun km, c phc dng li phn ln, v c th l mt "chimera" (tc c to nn t ha thch xng ca nhiu loi ng vt khc nhau). Hp s ca chng th gn nh rt ging vi mt loi Coelurosauria thu ban u[21].

Tin ha s khai ca chim

Chim pht trin a dng khc nhau trong sut thi k Phn trng[22]. Nhiu nhm vn gi nhng c im nguyn thy (symplesiomorphy), nh cnh c mng vut v c rng, d mt i mt cch c lp mt s nhm chim pha sau, bao gm chim hin i (Neornithes). Trong khi nhng dng ban u, nh Archaeopteryx v Jeholornis, vn mang nhng chic ui di v nhiu xng nh t tin chng[22], th chic ui ca nhng dng tin ha sau (nhnh Pygostylia) ngn hn vi kiu xng bnh li bn trong. Ging ln v a dng u tin m cc chim ui ngn tin ha l Enantiornithes, hay "chim o ngc", c mang tn ny bi kt cu xng vai ca chng o ngc li so vi cc loi chim hin i. Nhng Enantiornithes chim lnh a dng cc sinh thi, t nhng loi chim cao cng tm kim trong ct v n c cho ti nhng loi sng trn cy v n ht cy[22]. Nhiu loi tin ha sau cng thch ng vi vic n c, trong tiu biu nh phn lp Ichthyornithes ("chim c") c hnh dng ging mng bin[23]. B chim bini Trung sinh, Hesperornithiformes, cng tr nn thch nghi cao cho vic sn bt c trong mi trng bin, khi chng mt i kh nng bay v ch yu bi li trong nc. Mc d c tnh chuyn ha rt cao, Hesperornithiformes vn l i din tiu biu cho nhng h hng gn nht ca cc loi chim hin i[22].

Pht x chim hin i

Bao gm tt c cc loi chim hin i, phn lp Neornithes, da trn s khm ph chi Vegavis, hin nay c coi l tin ha t mt s ging c bn vo cui k Phn trng (Creta)[24], v sau tch thnh hai phn b Paleognathae v Neognathae. Paleognathae bao gm b Chim tinamou, sng ti khu vc Trung v Bc M, cng cc loi thuc b iu. nhm Neognathae cng c s phn ly c bn, to nn phn b Galloanserae, bao gm b Ngng v b G, cng phn b Neoaves gm cc loi cn li. Thi im bt u nhng qu trnh tch ra c tranh ci nhiu bi nh khoa hc. H ng vi nhau rng Neornithes tin ha trong k Phn trng, v s tch ra ca Galloanserae khi cc Neognathae khc c xy ra trc s kin tuyt chng K-T, tuy nhin c nhng kin khc nhau v vic s pht x tin ha ca nhm Neognathae din ra trc hay sau s tuyt chng ca cc loi khng long khc[25]. S bt ng ny mt phn bi s khc bit gia hai loi bng chng: phn tch v phn t

cho thy s pht x thuc v k Creta, nhng bng chng ha thch li ng h cho vic thuc v phn i Tam. Nhng c gng dung ha gia hai bng chng ha thch v phn t u dn n tranh ci[25][26]. Bn cnh , h thng phn loi chim cng l vn gy tranh lun. Sibley v Ahlquist trong cun Phylogeny and Classification of Birds (Pht sinh v phn loi chim) (1990) a ra mt cng trnh mang tnh bc ngot trong vic phn loi chim[27], mc d n thng xuyn b tranh ci v lin tc sa i. Hu ht cc bng chng cho thy dng nh s phn chia cc b ca lp Chim l chnh xc[28], nhng cc nh khoa hc vn khng thng nht c v mi quan h gia cc b vi nhau; cc bng chng t gii phu, ha thch v ADN hin i u c quy vo vn , nhng li khng c s ng thun no mnh. Nhng bng chng mi v ha thch v phn t gn y ang cung cp thm, gip r rng hn cho bc tranh v s tin ha ca cc b chim hin i.

Phn loi cc b chim hin i

Di y l danh sch cc b trong phn lp Chim hin i (Neornithes), s dng h thng phn loi truyn thng (thng gi l b Clements): Phn lp Neornithes Paleognathae:

1. Struthioniformes B iu 2. Tinamiformes Tinamou

Neognathae:

1. Anseriformes B Ngng 2. Galliformes B G 3. Charadriiformes B Choi choi (B R) 4. Gaviiformes Chim ln Gavia 5. Podicipediformes B Chim ln (B Le hi) 6. Procellariiformes B Hi u (B Chim bo bo) 7. Sphenisciformes B Chim cnh ct 8. Pelecaniformes B B nng 9. Phaethontiformes Chim nhit i (tropicbird) 10. Ciconiiformes B Hc 11. Cathartiformes Kn kn Tn Th gii 12. Phoenicopteriformes B Hng hc 13. Falconiformes B Ct 14. Gruiformes B Su 15. Pteroclidiformes - G ct (sandgrouse) 16. Columbiformes B B cu 17. Psittaciformes B Vt 18. Cuculiformes B Cu cu 19. Opisthocomiformes G mng Nam M (hoatzin) 20. Strigiformes B C 21. Caprimulgiformes B C mui 22. Apodiformes B Yn 23. Coraciiformes B S 24. Piciformes B G kin 25. Trogoniformes B Nuc 4

26. 27.

Coliiformes - Chim chut (mousebird) Passeriformes B S

H thng phn loi Sibley-Monroe c s khc bit c bn vi h thng trn, khi pht trin da trn cc d liu phn t. N cng c cng nhn tng i rng ri mt s mt, tiu biu gn y l vic s dng cc bng chng phn t, ha thch v gii phu trn phn b Galloanserae ng h cho h thng ny[25].

Phn b

Khu vc phn b loi s nh (Passer domesticus) m rng t ngt do cc hot ng ca con ngi[29].

Chim sng v sinh sn hu ht cc mi trng trn cn cng nh c by lc a, trong nm gi k lc phng Nam l loi hi u ptren tuyt(Pagodroma nivea) khi c th sinh sn khu vc ln su ti 440 kilmt trong chu Nam Cc[30]. Tnh a dng cao nht v cc loi chim thuc nhng khu vc nhit i. Trc y, ngi ta ngh rng tnh a dng cao l kt qu ca tc hnh thnh loi cao nhng khu vc ny, nhng nhng nghin cu gn y cho thy, khu vc c v cao, tc hnh thnh loi d cao hn nhng li b b tr bi tc tuyt chng m cng ln hn so vi vng nhit i[31]. Vi h chim li c cuc sng thch nghi c mi trng i dng, trong s , c nhng loi chim bin vo b ch duy sinh sn[32], v mt s chim cnh ct li c ghi nhn l c th ln su ti 300 mt (980 ft)[33]. Nhiu loi chim thnh lp nhng qun th giao phi nhng vng m chng c nhp ni bi con ngi. C nhng loi c nhp ni c ch , v d nh loi tr , c a i trn ton th gii nh mt loi chim sn bt[34]. S khc li mang tnh ngu nhin, mt v d l s thnh lp ca qun th vt thy tu ui di (Myiopsitta monachus) vi thnh ph ca Bc M, sau khi chng thot ra khi tnh trng nui nht[35]. Vi loi khc, bao gm c rui[36], diu vn u vng[37] v vt mo ngc hng (galah)[38] m rng phn b mt cch t nhin bn cnh khu vc phn b gc, bi cc hot ng nng nghip to nn sinh cnh mi thch hp cho chng.

Gii phu v sinh l

Gii phu bn ngoi ca mt con chim in hnh: 1 M, 2 u, 3 Mng mt, 4 ng t, 5 Lp lng cng cnh, 6 Lng bao cnh nh, 7 Lng vai, 8 Lng bao cnh gia, 9 Lng cnh b, 10 Phao cu, 11 Lng cnh s cp, 12 Hu mn, 13 i, 14 Khp c chy-chn, 15 Xng c chn, 16 Bn chn, 17 Xng chy, 18 Bng, 19 Sn, 20 c, 21 C, 22 Ym

Xt hnh thi bn ngoi, chim c c th dng hnh bu dc ngn, c di, u trn, nh. Ton thn chim c ph lng v, hai chi trc l cnh, hai chi sau c bn chn hnh tr gip chim i ng trn mt t[39]. So vi nhng ng vt c xng sng khc, c th chim c nhng c im thch nghi c bit, ch yu ph hp cho hot ng bay.

B xng v h c
Chim c b xng vi nhng chic xng nh v xp nhng chc[39]. Xng chim c nhng khoang cha y kh, c lin kt vi h h hp[40]. Xng s c ni lin v khng c cc ng khp s[41]. mt ln v c ngn cch bi mt vch xng. Ct sng ca chim c chia lm 4 phn: c, ngc, chu v ui, vi cc t sng c c tnh bin i cao v c bit linh hot; tuy nhin kh nng c ng ch cc t sng ngc tr i ch khng c cc t sau[42]. Nhng t cui kt hp vi khung chu to thnh khi xng cng[41], lm ch da vng chc cho cc

chi sau[39]. Tr nhng loi khng bit bay, xng sn chim dt v xng c pht trin c g li hi nhm gn kt vi cc c vn ng bay. Hai chi trc ca chim bin i thnh cnh[43]. Chim c tng cng khong 175 c trn c th[44], trong cc c pht trin lin quan n nhng hot ng nh bay (c vn ng cnh), chy (c i v c ng chn cc chim chy), x lng (c da) hay c ng u (c c)[39]. C ln nht l c ngc (c h cnh) c th chim t 15-20% khi lng c th con chim. Bn chn ca chim khng c c, tuy nhin li c cc gn c th t ng sit cht khp qup cc ngn chn quanh cnh cy khi chim u, gip chng bm lu v chc trn cnh m khng b mi[39].

Bi tit v tiu ha
Ging nh b st, chim cn bn l loi bi tit axt uric (uricotelic): ct chim lc cc cht thi gc nit t mu v bi tit chng di dng axt uric thay v ur hay amoniac thng qua ng niu trong rut. Chim khng c bng i hay niu o m bn ngoi, nn nc tiu ca chim c thi ra i km vi phn to thnh cht thi na rn[45][46]. Tuy vy, vn c nhng loi nh chim rui li thi hu ht cc cht thi nit di dng amoniac[47]. Cc loi chim thng thi creatin, hn l creatinin nh ng vt c v[41]. Loi cht ny, cng nh cc cht thi khc ca qu trnh tiu ha, c thi ra t huyt ca chim[48][49]. Huyt chim l mt ng m a chc nng, t tng cc cht thi ra ngoi ti giao phi v trng. Ngoi ra, nhiu loi chim cng (a quay tr li ng ming) cc thc n cha tiu ha cn d (pellet)[50]. H tiu ha chim c c im c nht, vi b phn diu lu tr thc n v m, cha cc hn c chim nut, c kh nng nghin thc n thay th cho b rng chng khng c[51]. Nhiu loi chim thch nghi cao vi vic tiu ha nhanh gip cho hot ng bay[52]. Mt s loi chim cng thch nghi bng cch s dng protein t nhiu b phn ca c th. cung cp thm nng lng trong qu trnh di tr[53].

H h hp v tun hon
Chim c mt trong nhng h h hp phc tp nht ca tt c cc loi ng vt[41]. Bn cnh phi, chim cn c 9 ti kh, l cc vi kh qun xuyn qua phi to thnh, dung tch hn phi nhiu ln, c vai tr cha kh h hp cng nh lm nh c th v iu ha thn nhit[39]. Lc chim ht vo, 75% lng khng kh sch khng i qua phi m ti trc tip cc ti kh sau sau lp y cc khoang trong xng. 25% lng kh cn li i trc tip vo phi. Khi chim th ra, nhng lung kh c s dng i ra ngoi phi v nhng kh sch cha trong xng cng lc li i vo phi. Theo cch , phi ca chng lun c duy tr cung cp khng kh sch trong c khi th ra v ht vo[54]. C quan to m thanh ca chim l minh qun (syrinx), mt khoang c vi mt s mng nh, t v tr im cui ca kh qun, ni kh qun phn thnh hai ph qun[55]. Tim chim c bn ngn, cung ng mch ch phi tham gia vo vng tun hon ln (khng ging nh ng vt c v l cung ng mch ch tri)[41]. Tnh mch ch di nhn mu t cc chi thng qua mt h gnh thn. Tim chim p nhanh v nhp tim t l nghch vi khi lng c th. g, tim p khong 250 ln/pht, sn tc u en khi ng l 500 ln, khi hot ng ln ti 1000 ln/pht[39], ring vi loi chim rui c (Archilochus colubris), tim mi pht c th p 1200 ln (20 ln/giy)[56]. iu ny gip cho mu chim lu thng nhanh, gip vn chuyn nhanh xy v cht dinh dng cho cc c quan, cung cp kp thi nng lng bay v duy tr hot ng mc cao. Bn cnh , hng cu chim c nhn (khc vi cc loi th)[57], nhiu v li hai mt, hemoglobin lin kt vi xy v cacbonic yu nn vic gii phng cc kh ny din ra nhanh trong mu. y l l do v sao chim c thn nhit cao, vo khong t 38-45,5oC, tu mi loi[39].

iu ha thn nhit
Chim l ng vt mu nng, thn nhit n nh khng ph thuc vo nhit bn ngoi. L sinh vt c ch s trao i cht cao[58], thn nhit chim cao v thng bin i t 40-42oC[39]. Cc loi chim nh thng c thn nhit t n nh hn cc loi chim ln, thay i theo ma hay thm ch hng m[59], mt v d l hng tc nh (Troglodytes aedon) c bin thin thn nhit trong 24 gi ln ti 8oC[60]. chim s sinh, d cng c thn nhit cao (xp x khong

38oC nhiu loi), nhng li cha c c ch hng nhit nh chim trng thnh, nn chng phi da vo nhit c th b m v t gi m[59][61]. Bn cnh trao i cht, khi nhit mi trng xung thp, chim thng x lng ngn khng cho khng kh tip xc vi da, co cc mch mu da, hay i khi run tng nhit c th. Kh nng iu ha thn nhit l mt nguyn nhn gip cc loi chim c th phn b rng trn khp hnh tinh[39].

Thn kinh v gic quan

chim s ng xanh (Cyanistes caeruleus, tn c Parus caeruleus), con trng c mt m trn u phn chiu tia cc tm, c chng ph by trc con mi qua t th x lng pha gy[62]. B lng ca c mo chu Phi cho php loi ny ng ha vi mi trng xung quanh

H thn kinh ca chim ph thuc nhiu vo kch thc c th[41]. Phn pht trin nht ca no iu khin cc chc nng lin quan n hot ng bay, trong khi tiu no phi hp s c ng v i no th iu khin cc kiu tp tnh, tm ng, giao phi v lm t. a phn cc loi chim u c khu gic km, ngoi tr mt s trng hp ni bt nh kiwi[63], kn kn Tn Th gii[64] v b Chim bo bo (Procellariiformes)[65]. Ngc li, th gic ca chng thng pht trin mc cao. Cc loi chim nc c thy tinh th linh hot c bit, cho php thch nghi vi vic nhn trong nc ln khng kh[41]. Mt s loi trong mt cn c hai vng hong im (fovea). Cc loi chim c kh nng nhn bit bn loi ngun sng (tetrachromacy), c cc t bo hnh nn nhy cm vi tia cc tm bn cnh cc di mu , xanh lam v xanh l cy[66]. iu ny cho php chng quan st c cc tia cc tm (UV), c lin quan trong qu trnh ve vn. Nhiu loi chim c nhng b lng sc s trong di tia cc tm m mt ngi khng nhn thy; mt s loi m hai gii tnh tng nh ging nhau vi mt thng, nhng li khc nhau di tia cc tm khi phn chiu nhng vt m trn b lng ca chng. Tia cc tm cn c s dng trong vic kim n - mt s loi ct tm kim con mi l nhng loi gm nhm bng cch xc nh tia UV phn chiu t du nc tiu trn mt t[67]. M mt ca chim khng c s dng nhy mt, thay v th chng dng mt m mt th ba di chuyn theo chiu ngang, gi l mng nhy[68]. Mng nhy cng bao ph mt v c vai tr nh mt loi knh p trng nhiu loi chim sng nc[41]. Vng mc chim c mt h thng cung cp mu hnh qut gi l lc (pecten)[41]. Hu ht chim khng th di chuyn mt ca chng, tr mt s ngoi l, nh chim cc [69]. Nhng loi chim vi mt hai bn u c tm nhn rng, cn nhng loi c mt trc mt, nh c, c th gic hai mt (binocular vision) v c th c tnh c chiu su ca tm nhn[70]. Chim khng c tai ngoi nhng tai chng c bao ph bi lng v, d mt s chim, nh cc chi c Asio, Bubo v Otus, lng v to nn nhng bi m tng t nh tai. Tai trong c mt c tai, nhng khng c dng xon nh cc loi th c v[71]. Tai chim c th nghe c m thanh c tn s gn vi tai ngi nhng li hn ngi nhiu ln v kh nng phn bit cng m thanh. So vi cc ng vt xng sng cn khc, xc gic chim km pht trin[39].

Cht ha hc
Mt vi ging chim c kh nng s dng cht ha hc chng li k th; vi loi thuc b Chim bo bo c th phun ra mt loi du nhn kh chu ln nhng k gy hn[72] hay mt s loipitohui t New Guinea, li c mt loi neurotoxin (c t thn kinh) mnh nm trn da v lng[73].

Lng v
Lng v l c im duy nht ch c cc loi chim. N h tr cho vic bay, to ra mt lp cch ly gip iu ha thn nhit, ngoi ra cn c s dng ph by, ngy trang hay lm tn hiu[41]. C mt s loi lng v, c s dng cho tng mc ch. Lng v mc ln t lp biu b ca da, ch nhng vng c bit gi l vng lng (pterylae). Kiu phn b ca nhng vng lng (pterylosis) c ng dng trong phn loi v h thng hc. S sp xp v v ngoi ca lng v trn c th, gi l b lng v (plumage), c th khc nhau trong loi da trn tui, a v by n[74] v gii tnh[75]. Cc loi chim thay lng theo nh k; b lng thng thng ca mt con chim m thay sau ma sinh sn gi l b lng "khng sinh sn" (non-breeding) - hay theo thut ng Humphrey-Parkes l b lng "c bn" (basic); cn b lng khi sinh sn hay cc dng khc ca b lng c bn gi theo Humphrey-Parkes l b lng "lun phin" (alternate)[76]. Vic thay (rng) lng l thng nin hu ht cc loi chim, d c nhng loi thay hai ln trong nm, hoc cc loi chim sn mi ln c th ch thay lng mt ln duy nht cho vi nm. C cc kiu thay lng khc nhau gia cc loi. cc loi chim s, lng bay (lng trn cnh v ui gip cho vic bay) c thay th mt ln trong mt thi gian vi nhng chic lng v s cp trong cng s thay u tin. Sau khi lng s cp c thay th ln th nm hay th su, nhng chic lng bc ba ngoi cng bt u rng, sau n lng th cp v cui cng qu trnh ny li i ti nhng chic lng pha ngoi (rng lng ly tm). Nhng lng bao (tetrix) ln c rng ln ng thi vi lng s cp[77]. Mt s nh cc loi, nh vt v ngng, li c th rng tt c cc lng bay cng lc, khin chng tm thi khng bay c[78] Theo mt quy lut chung, lng ui li rng v thay khi u t nhng cp trong cng[77]. D th, vic rung lng ly tm cng tm thy cc loi thuc h Tr[79]. Bn cnh , s rng lng ly tm ui c s thay i cc loi chim g kin v h ui cng, khi khi u vi nhng cp lng trong cng th hai v kt thc vi cp lng gia, v vn gia mt cp lng vi chc nng leo tro[77][80].

Vt Lory (Eos bornea) ang ra lng. Chn vi ca ca mt con g ty

cc loi s, kiu thay lng chung l lng cnh s cp thay trc, lng th cp thay sau, cn ui th khi u vi nhng lng gia[81]. Trc khi xy t, a phn cc con chim mi u rng phn lng gn bng to thnh mt "vt p" (brood patch). Phn da y s c cung cp tt t cc mch mu v h tr chim trong vic p trng[82]. Lng v i hi c gi gn v chim thng dnh khong 9% thi gian mi ngy dnh cho vic ra hay chi lng[83]. Chim dng m chi i cc ht nh t bn ngoi, ng thi bi ln mt loi sp tit ra t tuyn phao cu; loi sp ny s gip gi gn tnh linh hot ca lng v l mt tc nhn chng vi sinh vt, ngn chn s pht trin ca nhng vi khun lm gim cht lng lng[84]. Ngoi ra cht ny cn gip lng khng thm nc v cung cp vitamin D (chaergosterol bin i thnh vitamin D di nh mt tri)[39]. Bn cnh , mt s chim cn c hnh ng ch kin (hay cn trng khc) ln b lng ca chng, nhm ly axt formic tit ra t kin, c vai tr tiu dit nhng sinh vt k sinh trn lng[85].

M, chn v vy
M l mt cu trc bn ngoi ca chim, c s dng cho vic n v nhiu mc ch khc. Trong m khng c rng, m bao gm 2 phn, phn hm trn c bao ph bi mt bao v sng (rhamphotheca), cu to t keratin.

Gia phn trn m c hai l mi, n thng vi h h hp. mt s loi, m v l mi c bao bi mt phn m mm, sp, c gi l da gc m (cere)[86]. Bn chn chim c bao ph bi vy v c cc ngn chn vi mng vut. a phn cc loi chim u c 4 ngn, c chia thnh 5 kiu sp xp(dactyly) chnh, bao gm: anisodactyl (ngn khng u), zygodactyl (ngn kiu chn tro), heterodactyl (khc ngn), syndactyl (dnh ngn) vpamprodactyl (ngn nhn)[87]. nhiu loi chim, chn c mng, v cng c cc loi: c na mng (semipalmate), c mng hon ton (totipalmate), c mng kiu chn vt v c thy[88]. M cng mng vut u khng ngng pht trin v kch c[86]. Kiu chn v m ca mi loi nhiu khi rt khc nhau, ph thuc vo li sng v thc n ca chng. g trng v mt s loi chim khc, chn c phn sng cng mc ln, c gi l ca. Ging cc b phn m, vut v ca, vy chim l sn phm ca da[39], c to thnh t cng mt loi keratin. Vy c tm thy ch yu ngn chn v bn chn, nhng cng c th xut hin mt c chn mt vi loi. Hu ht cc loi chim, vy khng che ph ng k, ngoi tr trng hp g kin v ui cng. Vy ca chim c cho l tng ng vi vy ca ng vt c v v b st[89].

c im sinh hc
Di chuyn
Bay
i bng c chu Phi (Haliaeetus vocifer) ang chao ling trn khng.

Phn ln chim u c th bay, iu ny to nn s khc bit gia chng v hu ht cc loi ng vt c xng sng khc. Bay l phng tin chnh di chuyn ca a s loi chim, s dng cho cc hot ng sinh sn, kim thc n hay chy trn k th. Chim c nhiu c im thch nghi cho vic bay, bao gm b xng khi lng nh, hai khi c vn ng cnh ln (c ngc v c qu trn - supracoracoideus), cng nh hai chi trc bin i thnh cnh, c vai tr tng t nh cnh my bay[41]. Hnh dng v kch thc cnh thng thng xc nh kiu bay ca mi loi; nhiu loi chim c kh nng phi hp gia nhng c p cnh mnh m v kiu chao ling i hi t nng lng. Chim rui l mt trng hp c bit ca lp Chim, khi chng c th bay ln vn ti ch bng cch p cnh 15-80 ln mt giy (ty mi loi), v c bit c th bay ngc pha sau, mt kh nng m khng nhm chim no khc c[90]. Chim ct Falco peregrinus vi si cnh rng l k nhanh nht trong gii ng vt, vi nhng c ling xung di t tc hn 322 km/h (200 mph)[91]. Khong 60 loi chim cn tn ti khng bit bay, cng nh vi nhiu loi chim tuyt chng[92]. Vic khng bit bay ch yu xut hin cc loi sng bit lp trn cc o, ph thuc vo cc ngun ti nguyn gii hn v khng c k sn mi[93]. D khng bit bay, nhng con chim cnh ct vn s dng c v cch di chuyn tng ng "bay" xuyn qua ln nc, tng t cc loi chim anca, ht nc hay chim bo bo[94].

Cc kiu di chuyn khc

G kin Hispaniola(Melanerpes striatus) dng mng bm ln cy. Chim cnh ct hong (Aptenodytes forsteri) lt mnh trn tuyt.

Bn cnh bay, cc loi chim cn c cc kiu di chuyn khc nh leo tro, bi ln hay i trn mt t[39]. Leo tro l kiu di chuyn nguyn thy ca nhiu loi chim tin s, v d nh Archaeopteryx s dng mng vut leo ln cy sau th mnh lt xung t[95]. Tri qua thi gian, chn ca cc loi chim leo tro nguyn thy (vi 3 ngn pha trc, 1 ngn pha sau) c bin i thnh chn ca cc chim leo tro hin i[39], vi mng khe cng 2 ngn trc v 2 ngn sau (kiu ngn zygodactyly). S loi chim leo tro cn tn ti khng nhiu, bao gm cc loi thuc cc h G kin,ui cng, Tro cy cng nh vt[96] v mt s loi khc. Cc loi ny u c cch leo tro ring ca mnh, mt s loi nh vt s dng m nh mt chn th ba tro cy[97]. Chim g kin dng mng sc bm v dng ui cng nh mt im ta, di chuyn trn cc thn cy[98]. Cc loi chim sng gn nc c cc kiu di chuyn khc nhau. Nhng loi bi ln gii c nhng c im thch nghi nh chn c mng hay b lng khng thm nc[99]. Chng c chia lm 2 loi da trn cch thc bi[100]: nh chn, tc dng chn nh mt mi cho to sc y, v d chim cc, chim ln, gavia v h Vt, v nh cnh - ch yu l cc loi sng bin, tiu biu l chim cnh ct, chim anca hay hi u ptren ln. Cc loi chim ln nh cnh nhn chung thng nhanh hn cc loi s dng chn[101]. Tuy nhin, d dng chn hay dng cnh th u khin cc loi ny b hn ch cc cch di chuyn khc, nh bay hay di chuyn trn mt t. Cc loi chim cnh ct khng bit bay c th l sinh vt thch nghi vi nc nht trong s cc loi chim bi ln[99], c bit chim cnh ct Gentoo(Pygoscelis papua) l loi chim bi nhanh nht, vi tc t 36 km/h (22 mph)[102]. Bn cnh , cn c cc loi ln bng cch lao thng t trn cao xuyn qua ln nc bt mi, v d chim in, bin, b nng nu v mt s nhn bin. Nhng loi chim nh hng hc, su, dic, c... v nh hn nh d hay choi choi l nhng loi chim li nc, vi i chn di, mnh, c th i qua nc d dng m c th khng b t, dng chn hay m kim thc n[99]. Hu ht cc loi chim u c th di chuyn trn mt t qua hai cch: i v chy[103], vi kh nng khc nhau mi loi. Tuy nhin, cng c nhng ngoi l nh chim yn, khi khng c kh nng u hay chy do i chn qu yu t, m ch c th dng chn bm ln cc b mt thng; hay hu ht cc loichim ln gavia cng khng th i ng bnh thng trn mt t do kt cu i chn ch ph hp vi vic bi ln[104]. Ngc li, c nhng chim nh b iu, tinamou, tc, g maleo cng nh nhiu loi thuc b G[103], pht trin kt cu chn thch hp, l i din ca nhng loi chim di chuyn tt trn mt t. Cc loi chim nc thng di chuyn trn mt t rt kh khn; d

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th, di chuyn nhanh hn, chim cnh ct li c mt phng thc c bit, l "lt vn" trn phn bng ca chng, s dng cnh v chn y thn mnh lt i trn bng tuyt[105].

Di truyn
Chim c hai gii tnh: trng (c) v mi (ci). Gii tnh ca chim c xc nh bi hai nhim sc th gii tnh Z v W, khng ging cp nhim sc th (NST) X v Y ng vt c v. Cc con trng c hai nhim sc th Z (ZZ) v con mi c mt NST Z v mt NST W (WZ)[41]. Gn nh ton b cc loi chim, gii tnh mt c th c xc nh ti thi im th tinh. Tuy nhin, mt nghin cu gn y cho thy s xc nh gii tnh ph thuc nhit nhng con g ty bi rm c (Alectura lathami), khi m nhit cng cao trong qu trnh p trng th s to t l mi trn trng cao hn[106].

Sc khe
Tui th
Tui th ca chim rt khc nhau ty theo loi, c th t ba n bn nm tui i vi mt s loi chim s v hn 50 nm i vi mt s loi hi u, v thm ch hn 60 nm tui i vi mt s loi him c nh chim kakapo (Strigops habroptilus). Trong cng mt loi chim, vic thay lng khng nhng ph thuc vo ma, m cn vo tui ca chim, do vy vic bit r cc thng tin ny s gip tnh ra tui ca nhiu loi chim hoang d[107]. Ngoi ra mc kh ha khung xng cng l mt c im c s dng c tnh ra tui ca chim.

K sinh
Cc sinh vt k sinh ph bin chim ch yu thuc v cc nhm gh, rn v chy. Cc loi k sinh khc nh hn nh mt s loi thuc ng vt nguyn sinh, thng gy bnh cho chim.

Dermanyssus gallinae, mt con rn k sinh g. Mt trng hp vim phi nng do nhimChlamydia (bnh virus vt) mt con dicEgretta caerulea

t nht 2.500 loi gh thuc 40 h[108] sng gn vi cc loi chim, t t chim, lng, thm ch m chim nh mt s loi gh k sinh trn chim rui. Cc loi gh ny c th ch n thun l k sinh v hi hoc c th gy nga cho ng vt ch, thm ch l k sinh c hi nh cc loi thuc chi Dermanyssus vOrnithonyssus. Tt c cc loi chim u b nh hng bi sinh vt k sinh, ngay c chim cnh ct[108] cng c ve k sinh trn c th. i sng cc loi ve tt nhin ph thuc vo tng ging chim c th; tuy th, u trng ve thng sng trong t chim. Cc loi k sinh ny c chu k sinh sn ngn v c th sinh si rt nhanh chng. Mt s loi k sinh sng bng lp da cht ca chim, s khc nh cc loi k sinh chim rui th c mang t bng hoa ny sang bng khoa khc v sng bng mt hoa. Trong cc t chim, ngi ta thm ch cn pht hin thy cc loi k sinh nh sng bm vo cc loi k sinh ln hn[108]. Mt s ln cc loi k sinh c th gy hi cho chim non hoc g con[108]. Tuy th, mt s nghin cu cho thy s hi sinh ny khng phi ch c hi cho chim[108]. C kh nhiu nghin cu v cc tng tc phc tp gia sinh vt k sinh v chim, v nhiu yu t c tm hiu. V cc nghin cu cho thy vic a ra cc quy tc n gin trong trng hp ny l khng d dng.

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Mt s loi thuc b n lng (nh Ischnocera) thng sng bm mt s loi chim nht nh. Rt nhiu loi thuc ngnh Giun dp nh sn dy hay sn l c th ly nhim cc loi chim v cc con chim ny c th mang chng i t lc a ny sang lc a khc. Chng hn nh cc loi chim bin khi n cc loi nhuyn th thuc chiCerastoderma, c th pht tn cc loi sn l k sinh (cc chi Meiogymnophalus, Himasthla,...), truyn sang cc ng vt ch khc nh cc loi chim hoc cc loi ng vt thn mm khc[109].

Bnh tt
Chim v gia cm l mt trong nhng vc t truyn bnh ch yu cc c ly xa i vi con ngi nh cc bnh virt vt, bnh vi khun Salmonella,bnh vi khun xon (do Campylobacter), bnh lao gia cm (do Mycobacterium), cm gia cm, st hi ly (bnh do loi k sinh Giardia lamblia, v bnh Cryptosporidium[110]. Cc bnh ny cng nh s ly lan ca chng cng c nghin cu rt k. Cng do tm ph bin ca ngnh nui gia cm, vic pht hin ra cc bnh gia cm c th khin chnh quyn a phng p dng cc bin php trit v ngt ngho i vi vic chn nui ny. Vo thng 9 nm 2007, 205.000 con gia cm ti Bavire b thiu hy[111], 160.000 gia cm ti Bangladesh cng b thiu hy vo thng 2 nm 2008[112] do pht hin ra dch cm gia cm... Mt s bnh c th nh bnh Pacheco ch c cc loi thuc b Vt.

Cc bnh khc St rt gia cm Bnh Newcastle Bnh do vi khun thuc lp th cp)

c im chung ca Lp Chim (Aves)

Bnh m v Tiu chy ghay gia cm Vim kh qun chim lng(PBFD) u ma gia cm(dng Gin d dy bch cu) (PSD, thng b Vt)

Bnh nhim nm Candida gia cm Bnh nm qut

Mollicutes (vi khun khng c mng Bnh Gumboro

Lp Chim c s lng loi phong ph, c khong 8.600 loi, phn b khp mi min trn Tri t. Trn sut 130 triu nm tin ho theo hng thch nghi vi chuyn vn bay nn tt c cc loi chim hin i t chim rui ch nng 1,8g n iu chu Phi to ln nng gn 80kg u c cu trc c th ng dng. Hnh thi v cu to c th chim c c im sau: - C th chim c hnh dng van ngn, chia bn phn: u, c, thn v ui. Ton thn ph lng v. Chi trc thng bin i thnh cnh thch nghi bay. Chi sau bin i khc nhau thch hp vi u trn cnh cy, i trn mt t v bi trong nc. Bn chn 4 ngn. - Da mng, hu nh khng c tuyn, tr tuyn phao cu ton thn ph lng v, mt iu kin rt cn thit cho chim c th bay c. Chn ph vy sng. - B xng hon ton bng xng. Tuy nhin thch nghi vi s bay, xng c cu to xp, nhiu khoang kh. Hp s ln, c mt li cu chm, xng hm khng c rng ch ph m sng. Cc t sng thn c xu hng gn li vi nhau, trong khi cc t sng c li khp vi nhau rt linh hot. xng sn nh, xng c pht trin to nn g li hi. ai vai v xng chi trc bin i thch nghi vi s bay. ai hng c cu to thch nghi vi vic trng ln c v cng. - H thn kinh pht trin cao: Bn cu no, thu th gic v tiu no ln, thu khu gic nh. No b un khc r rng. C 12 i dy thn kinh no. - Gic quan pht trin: C quan thnh gic gm tai trong, gia v ngoi, c vnh tai n gin. C quan th gic pht trin, l b phn nh hng khi bay. Khu gic km pht trin. - H tun hon kh pht trin: Tim 4 ngn, ch cn cung ch ng mch phi. H mch mu gan thn tiu gim. C 2 vng tun hon cch bit, mu khng pha trn, t bo mu c nhn. - H hp bng phi, c h ti kh pht trin len li trong ni quan, da v xng. H thng ti kh gip chim gim nh trng lng, cch nhit v c bit l tham gia h hp khi chim bay. - C quan tiu ho bin i quan trng nh khng c rng, thiu rut thng tch tr phn, cc phn ni quan u tp trung v pha trc c th. - H bi tit l hu thn. ng dn niu ni vi huyt, khng c bng i, nc tiu c, sn phm bi tit ging nh b st l axit uric, c thi ra cng vi phn. - H SD phn tnh. Con c c i tinh hon khng bng nhau, tinh qun vo huyt, c quan giao cu ch c vt ngan, chim chy... Con ci ch c 1 bung trng v mt ng dn trng tri, do vy trng 12

lng c th chim gim i nhiu. - Th tinh trong, p trng v chm sc con. Trng nhiu non hong, c v mng trong v v vi ngoi. Pht trin c hnh thnh mng phi. Chim non mi n thng l chim khe mnh.

Y Hc C Truyn Quan Nim V T V

Trng thi sinh l, bnh l ng tiu ha t xa xa c y hc c truyn (YHCT) lu tm nghin cu. Trong h thng cu trc - chc nng (ng tng lc ph) ca c th m YHCT gi l tng tng, t v l 2 c quan c nhiu im gn gi vi h thng ng tiu ha ca y hc hin i (YHH). V c sch c m t l mt c quan rng, trn tip vi thc qun, di thng vi tiu trng, ming trn gi l "b mn", ming di gi l "u mn", b mn cn gi l "thng qun", u mn cn gi l "h qun", c ba vng gi l "v qun". Thc n t ming qua thc qun ri vo v, c v lm chn nh, cho nn v gi l "i thng", l ci kho ln, ci "b cha n". T l mt c quan c nm bn tri ca v c chc nng hp thu v vn chuyn cht dinh dng, YHCT gi l c cng nng vn ha. T v v hp tc vi nhau hon thnh chc nng tiu ha, hp thu thc n v chuyn vn cht dinh dng. T v c quy np theo hnh "Th" trong h thng ng hnh (mc, ha, th, kim, thy) ca trit hc c i phng ng. "Th" l m ra vn vt cng ging nh t v c chc nng hp thu, chuyn vn cht dinh dng nui cc b phn khc trong c th. V vy, vai tr ca t v trong h thng tng tng ca YHCT l c bit quan trng.

Tht vy, thn, t (lch) l nhng b phn quan trng ca h tiu ha c tc ng n tr no v cho rng, bnh pht sinh ch yu l do tm t suy nhc, thn tinh h yu gy nn. Bi ng y quan nim tm t ch huyt, thn ch tinh ty, l tinh hoa ca ng tng. Thn l tng ca thy ha, chn m v chn dng u . Thn m v thn dng tuy l hai mt i lp nhau nhng lun lun c s h tr qua li ln nhau nhm duy tr s tn ti v chc nng ca thn...

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