M hnh ABC v s pht trin ca hoa

Hoa Arabidopsis thaliana

C khong 350000 loi thc vt ht kn khc nhau, v hu ht chng c phn loi da trn cc c im ca
hoa. Mt im m hoa ca cc loi thc vt ging, m cng khc nhau nht l kiu dng ca cc b phn ca hoa.
Hu ht cc loi hoa bao gm 4 c quan sp xp theo vng ng tm: l i (sepals), cnh hoa (petals), nh hoa
(stamens), v l non (carpels). Nh hoa v l non c tt c cc loi ht kn, tt nhin l m bo kh nng
sinh sn v duy tr ni ging. Ngc li, khng phi loi no cng c l i v cnh hoa ring r. Trong trng hp
nh th, hoa ch c 3 c quan xp theo vng ng tm l tepals (o ch ca petals), stamens v carpels.

Hnh 1. (A) Cc b phn ca hoa Arabidopsis. (B) Hoa ca cy l l (Hellebore) khng c cnh hoa
tht: cu trc ging cnh hoa (v i hoa) gi l tepal.

Hnh 2. a dng v cu trc hoa ca thc vt. (A) Hoa ca loi Passiflora (cy chanh leo) c tn hoa
hnh si nm gia cnh hoa v nh/nhy. (B) Echinopsis backebergii, hoa ca loi n c mt phn nh
hp vi vi nhy to thnh mt cu trc mi gi l gynostegium. (C) Iris pseudoacorus (hoa Din v).
Mi tn cho thy vi nhy hnh cnh hoa. (D) Echinopsis backebergii (thuc h xng rng) c i
hoa v cnh hoa khng phn bit r rng (tepal). (E) Paphiopedilum spicerianum (h phong lan).
Cnh gia (mi tn) c cho l pht trin t tepal v nh hoa. Ngoi ra, cc b phn sinh sn dung
hp li thnh mt cu trc duy nht (u mi tn). (F) Aristolochia fimbriata (h Mc hng nam) c
i hoa hnh ng.
Gii thch c s bo ton cng nh s a dng trong cu trc hoa ca thc vt ht kn l mt vn cc k th
v. lm c iu ny, v tng t nh cc nghin cu khc, cc nh khoa hc tp trung phn tch cc t
bin v cu trc hoa trn cc cy m hnh, tiu biu nht l trn cy ci du Arabidopsis thaliana v cy mm
ch Antirrhinum majus (snapdragon). Cc nghin cu dn n mt m hnh tng i n gin v v cng tinh
t gii thch cho s pht trin ca cu trc hoa, gi l m hnh ABC.

Vic thu thp cc t bin v kiu hnh hoa c tin hnh t lu, nhng cc nghin cu h thng mc
phn t ch c bt u t cui nhng nm 1980. Nhn chung, cc t bin hoa c phn chia thnh 3 nhm, A,
B v C. Cc t bin nhm A c l non vng ngoi cng v nh vng th 2; t bin nhm B c l i vng
th 2 v l non vng th 3; v t bin nhm C c cnh hoa vng th 3 v l i vng th 4. B phn hoa
cc vng khc khng thay i.

Hnh 3. Ba nhm t bin nh hng n cu trc ca hoa Arabidopsis. (A) Hoa bnh thng (kiu
di). (B) t bin nhm A. (C) t bin nhm B. (D) t bin nhm C.

T cc c th t bin cc nh khoa hc phn lp c nhiu gene lin quan; cc gene ny c c im chung

l t bin trn mt gene ch nh hng n s pht trin ca 2 vng hoa lin k, khng phi ton b hoa. Nm
1991 Bowman v cng s xut m hnh ABC gii thch cho iu ny. Theo , s tng tc kt hp ca cc
gene thuc 3 nhm gene quy nh s hnh thnh cc vng hoa ( hoa, cnh hoa, nh v l non) b nh hng
v sinh ra cc t bin ny. Cc gene cng c phn nhm tng t nh cc t bin l A, B v C, trong
: gene thuc nhm A hnh thnh i hoa, A + B hnh thnh cnh hoa, B + C hnh thnh nh hoa v C hnh thnh
l non.

Hnh 4. M hnh ABC gii thch s hnh thnh cc vng hoa, na trn l mt bng hoa Arabidopsis
cha pht trin y c t mu tng ng vi s biu hin ca cc gene. Gene nhm A quy nh
s hnh thnh i hoa; Gene nhm A tng tc vi nhm B (mi tn mu v vng mu xanh
dng) quy nh s hnh thnh cnh hoa; gene nhm B tng tc vi gene nhm C (mi tn mu
v vng mu cam) quy nh s hnh thnh nh; v cui cng, gene thuc nhm C quy nh s hnh
thnh l non. Cc gene nhm A v C c tc ng i khng nhau.

C th dng m hnh ABC gii thch cc dng t bin m t trn nh sau:

Hnh 5: M hnh ABC gii thch cho cc th t bin. cy kiu hnh di, cc gene nhm A, B, C hot
ng nhp nhng vi nhau to thnh hoa y . t bin nhm A, gene nhm A b bt hot dn
n vic gene nhm C biu hin vng ngoi cng vn d l vng biu hin ca gene nhm A. Do ,
l non xut hin vng ngoi cng. Vng th 2 v th 3 gi y ch c s tng tc ca gene nhm
B v nhm C (khng cn tng tc gia nhm B v A) nn xut hin nh. t bin nhm B, cc
gene nhm B b bt hot, gi ch cn gene gia nhm A v C hot ng i khng nhau. Gene nhm
A biu hin 2 vng ngoi to ra l i, cn gene nhm C biu hin 2 vng trong to ra non.
t bin nhm C, cc gene nhm C b bt hot dn n vic gene nhm A biu hin vng trong
cng vn d l vng biu hin ca gene nhm C. Do , l i xut hin vng trong cng. Vng th
2 v th 3 gi y ch c s tng tc ca gene nhm B v nhm A (khng cn tng tc gia nhm
B v C) nn xut hin cnh hoa.
Qua phn tch cc nhm t bin trn Arabidopsis v hoa snapdragon, cc gene thuc tng nhm A, B, C c
pht hin. Ngoi tr gene APETALA2, tt c cc gene cn li u m ha cho cc yu t phin m, l cc phn t
protein c kh nng nhn bit v gn kt vi mt on DNA c hiu (thng nm v tr ca promoter) v t
nh hng n s biu hin ca gene lin quan.

Hnh 6. Cc gene thuc nhm A, B v C pht hin trn Arabidopsis v Snapdragon.

M hnh ABC hp dn bi tnh n gin nhng hiu qu ca n. iu ny c th hin thng qua kh nng d
on cao ca m hnh khi d on kiu hnh ca hoa t bin kp. V d, theo nh m hnh th nu cy b t bin
gene nhm A v B th ch cn gene nhm C hot ng, nh th hoa s ch c l non. V d na, nu cy b t
bin gene nhm B v C th ch cn gene nhm A hot ng, nh th hoa s ch c l i. Kt qu thc nghim
hon ton ph hp vi m hnh.

Hnh 7. Kiu hnh hoa ca cy Arabidopsis t bin kp. (A) t bin gene nhm A (apelata2) v B
(pisillata-1). (B) t bin gene nhm A (apelata2) v C (agamous-1). (C) t bin gene nhm B
(pisillata-1) v C (agamous-1). v (D) t bin gene nhm B (apelata3) v C (agamous-1).
Th cn trong trng hp t bin b ba th sao, nu gene thuc c ba nhm u b bt hot? Trong trng hp
ny s khng c b phn no ca hoa c hnh thnh, thay vo hoa s tr thnh mt c quan ging nh l.

Hnh 8. t bin b ba gene thuc c ba nhm A, B v C bin hoa ca Arabidopsis thnh c quan vi nhng c trng ca l. (A) B ba
gene apetala2 agamous-1 apetala3. (B) B ba gene apetala2 agamous-1 pistillata-1. (C) apetala2 agamous-1 pistillata-1.

Nh vy, cc gene nhm A, B, C c chng minh l cn thit trong s pht trin v hnh thnh cc vng hoa.
Cu hi by gi l, liu cc gene cha, hay vn cn nhng gene khc tc ng n tin trnh ny m

cha c pht hin ra. Nu qu thc 3 nhm gene ny l cn v ri, th vic biu hin sai v tr cc gene ny
trn l s bin l thnh c quan ging hoa. Tuy nhin, cc th nghim cho thy ch 3 nhm gene ny l khng
bin l thnh hoa.

T , Pelaz v cng s suy lun rng cn c nhng gene khc na hot ng cng vi cc gene A, B, C pht
hin. Bng tm kim trnh t tng ng trn genome, Pelaz v cs tm thm c cc gene mi, t tn l
SEPALLATA1, SEPALLATA2 v SEPALLATA3, l do t bin trn c 3 gene ny lm cho hoa ch gm ton l i
(sepals) (t bin ch mt gene khng thay i kiu hnh nhiu, cho thy 3 gene ny c tc dng b tr ln nhau).
Kiu hnh ny ging nh trong trng hp t bin kp gene nhm B v C. Suy ra, hot ng ca gene
SEPALLATA1,2,3 l cn thit cho hot ng ca gene nhm B v C.

Hnh 9. t bin b ba sep1 sep2 sep3 cho kiu hnh hoa ton l i vng 1, 2 v 3, cn vng 4 l mt bng hoa mi vi 3 vng ngoi cng
ton l i cn vng 4 l mt bng hoa mi. Nh vy kiu hnh hoa ny l lp li v tn. (a) Hoa kiu di. (b) t bin b ba sep1 sep2 sep3. (c)
Tch ri cc vng, ch vng 4 l mt bng hoa mi. (d) Kiu hoa t bin kp gene nhm B v C.

Tip tc nghin cu, Pelaz v cng s chng minh rng cc gene nhm A, B, C v gene SEPALLATA l bin
i l sang hoa. Nhm thu c kt qu thuyt phc khi bin ton b l cy Arabidopsis thnh cu trc ging
nh cnh hoa.

Hnh 10. Biu hin cc gene SEP (SEP3), nhm A (AP1) v nhm B (AP3, PI) lm ton b cy Arabidopsis chuyn thnh cu trc ging mt bng
hoa ln.

Nm 2004, Gary Ditta v cng s tm ra thm c mt gene SEPALLATA na l SEPALLATA4. t bin trn gene
khng lm cho cu trc hoa khc vi kiu di bao nhiu, chng t hot ng b tr ca 4 gene SEP. Tuy nhin, khi
bt hot c 4 gene SEP1,2,3,4 trn Arabidopsis th mang t bin sinh ra cc hoa vi nhng b phn c cu trc
ging nh l. Lu l, trc Pelaz v cng s lm t bin b ba gene SEP1,2,3 th lm cho hoa ch ton gm
l i (tng t t bin kp gene nhm B v C). Kt hp li, ta c th nhn xt l gene SEPALLATA4 l cn thit
cho s hot ng ca geen nhm A.

Hnh 11. Cu trc hoa ca cc dng t bin gene SEP so vi kiu di. (E) v (H) Hoa kiu di. (F v I) t bin b ba, hoa ch ton l i. (G) v
(J) t bin b bn, cc b phn trn hoa trng ging nh l. Ch cc mi tn vng ch trichoma (lng l).

T s khm ph ra chc nng quan trng ca cc gene SEPALLATA trn ton b hoa m cc bin th ca m hnh
ABC truyn thng ra i, nh l m hnh ABCE v m hnh quartet (b t). Trong cc m hnh ny, cc gene SEP
c xp vo nhm E.

Hnh 12. M hnh ABCE xut hin sau s khm ph ra 4 gene SEPALLATA. Cc gene SEP (nhm E) c vai tr tt c cc vng hoa.

Hnh 12. M hnh quartet. Cc hnh trn tng trng cho cc transcription factor m ha bi cc gene nhm A (AP1), B (AP3 v PI), C (AG) v E
(SEP). S pht trin ca mt vng hoa l do s kt hp ca cc transcription factor ny quy nh.

Tm li, m hnh ABC (v cc bin th ca n) tuy n gin nhng kh mnh m trong vic l gii s pht trin
cu trc ca hoa. Chnh v th n vn c s dng rng ri k t khi c xut (nm 1991) cho n nay.

Ti liu tham kho (click vo tn bi bo download)

1. Bowman, J. L., Smyth, D. R., & Meyerowitz, E. M. (1991). Genetic interactions among floral
homeotic genes of Arabidopsis. Development (Cambridge, England), 112(1), 120. Retrieved from
http://www.ncbi.nlm.nih.gov/pubmed/1685111

2. Bowman, J. L., Smyth, D. R., & Meyerowitz, E. M. (2012). The ABC model of flower development:
then and now. Development (Cambridge, England), 139(22), 40958. doi:10.1242/dev.083972

3. Causier, B., Schwarz-Sommer, Z., & Davies, B. (2010). Floral organ identity: 20 years of ABCs.
Seminars in cell & developmental biology, 21(1), 739. doi:10.1016/j.semcdb.2009.10.005

4. Theissen, G., & Saedler, H. (2001). Plant biology. Floral quartets. Nature, 409(6819), 46971.
doi:10.1038/35054172

5. Coen, E. S., & Meyerowitz, E. M. (1991). The war of the whorls: genetic interactions controlling
flower development. Nature, 353(6339), 317. doi:10.1038/353031a0

6. Ditta, G., Pinyopich, A., Robles, P., Pelaz, S., & Yanofsky, M. F. (2004). The SEP4 gene of Arabidopsis
thaliana functions in floral organ and meristem identity. Current biology : CB, 14(21), 193540.
doi:10.1016/j.cub.2004.10.028

7. Litt, A., & Kramer, E. M. (2010). The ABC model and the diversification of floral organ identity.
Seminars in cell & developmental biology, 21(1), 12937. doi:10.1016/j.semcdb.2009.11.019

8. Pelaz, S., Tapia-Lpez, R., Alvarez-Buylla, E. R., & Yanofsky, M. F. (2001). Conversion of leaves into
petals in Arabidopsis. Current biology : CB, 11(3), 1824. doi:10.1016/s0960-9822(01)00024-0

9. Soltis, D. E., Chanderbali, A. S., Kim, S., Buzgo, M., & Soltis, P. S. (2007). The ABC model and its
applicability to basal angiosperms. Annals of botany, 100(2), 15563. doi:10.1093/aob/mcm117