Jose Hernandes PDF

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José Hernandes Lopes Filho
Desenvolvimento da folha e axila foliar em

Portulacaceae e Talinaceae: homologias primárias
no clado ACPT.
Leaf and leaf axil development in Portulacaceae

and Talinaceae: primary homologies in ACPT
clade.
São Paulo
2010
José Hernandes Lopes Filho
Desenvolvimento da folha e axila foliar em

Portulacaceae e Talinaceae: homologias
primárias no clado ACPT.
Leaf and leaf axil development in

Portulacaceae and Talinaceae: primary
homologies in ACPT clade.
Dissertação apresentada ao Instituto

de Biociências da Universidade de
São Paulo para a obtenção de Título
de Mestre em Botânica, na Área de
Anatomia Vegetal.
Orientadora: Profa. Dra. Gladys

Flávia de A. Melo-de-Pinna
São Paulo
2010
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Comissão Julgadora:
________________________ _______________________
Prof(a). Dr(a). Prof(a). Dr(a).
______________________________
Profa. Dra. Gladys Flávia de A. Melo de Pinna
Orientadora
Aos meus avós, pais e irmãs,
por todo o carinho, apoio e
compreensão.
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2D2PD24A426DA3D2D !D562"D7D769D526DAD42752D2D25D5D4D
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37
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1
2
3 4
5 6
Figura 1. Aspecto geral das espécies de Portulaca e Talinum estudadas. A. P. werdermannii.

B. P. hirsutissima. C. P. hirsutissima. D. P. mucronata. E. P. halimoides. E. T. paniculatum.
1 2
3 4
Figura 2. Se7ões longitudinais da região nodal de esp8cies de Portulaca e Talinum. Notar a

constri7ão abaxial (setas) presente em todas as esp8cies. A. P. hirsutissima, notar sinuosidade
do caule dada pela expansão da base foliar. B. P. halimoides. C. P. mucronata. D. P.
werdermannii. E. T. paniculatum. Escala = 4009m (A), 2009m (B-E).
1 2
3 4
Figura 3. Se7ões longitudinais da axila foliar de esp8cies de Portulaca exibindo região basal
meristemAtica dos tricomas. A. P. halimoides. B. P. hirsutissima. C. P. werdermannii. D. P.
mucronata. Notar rApido alongamento que ocorre em c8lulas que se distanciam da região
basal. Escala = 1009m (A-B, D), 509m (C).
1 2
3 4
Figura 4. Seções transversais (A-E), longitudinal (F) e microscopia eletrônica de varredura (G), da região
da axila foliar de Portulaca. A. P. halimoides. B. P. mucronata. C. P. hirsutissima. D. P. mucronata. E. P.
mucronata. F. P. halimoides. G. P. hamimoides (notar paredes externas do tricoma colapsadas, enquanto
paredes internas permanecem rígidas). Escala = 50µm (D-E, G), 100µm (A-C, F).
1 2
3 4
Figura 5. Se7ões longitudinais (A, B e D) e transversal (C) da região da axila foliar de

Portulaca, mostrando a distribui7ão dos tricomas. A. P. werdermannii. B. P. hirsutissima. C. P.
halimoides. Notar tricomas nas laterais da inser7ão da folha (figura 3C). D. P. werdermannii.
Notar tricomas na região da constri7ão abaxial. Escala = 1009m (A, C-D), 2009m (B).
1 2
3 4
5 6
Figura 6. Se7ões transversais (A, B, D e E) e longitudinais (C e F) da região da axila foliar de

Portulaca, exibindo a instala7ão de felogênio. A-B. P. hirsutissima. Notar divisões celulares na
epiderme e camada subepid8rmica. C. P. halimoides. D-E. P. hirsutissima. F. P. werdermannii.
Escala = 2009m (A), 1009m (B-D, F), 509m (E).
1 2
3 4
5 6
Figura 7. Se7ões transversais (A-D) e longitudinais (E e F) da região da axila foliar de

Portulaca, mostrando a distribui7ão da periderme. A. P. halimoides. B. P. hirsutissima. C. P.
halimoides. D. P. werdermannii. E-F. P. werdermannii. Notar que em diferentes regiões da
constri7ão pode estar presente apenas tricomas, ou tamb8m periderme. Escala = 4009m (B),
2009m (A, C-F).
1 2
3 4
Figura 8. Se7ões transversais (A-D) e longitudinal (E) da região da axila foliar de Portulaca.
A. P. hirsutissima. Notar cristais na camada mais interna da periderme sob luz polarizada. B. P.
hirsutissima. Imagem sob luz polarizada. C. P. hirsutissima. Mesma imagem de B sem luz
polarizada. D. P. werdermannii. E. P. mucronata. Notar gema axilar com forma7ão de
primCrdios foliares e ausência da periderme. Escala = 1009m (A), 2009m (B-E).
1
1 2
1
2
1
1
1
3 4
1
2 1
2
1
5 6 1
1
1
2
2
1
B 1 D
Figura 9. Se7ões transversais da região nodal de Talinum paniculatum. Seqüência de
diferencia7ão das estruturas formadas a partir da gema axilar. 1 = par de profilos; 2 = estrutura
abaxial. Escala = 3009m (A), 2009m (F, H), 1009m (B-E, G).
1 2
3 4
5 6
Figura 10. Se7ões longitudinais (A-D) e transversais (E, F) da região da axila foliar de
Talinum paniculatum mostrando profilos (seta) e estrutura abaxial (ponta de seta). Notar
ausência de vascularização e presença de cavidades secretoras na região apical das estruturas
(E e F). Escala = 2009m (A, C-D), 1009m (B, E), 509m (F).
1 2
3 4
* *
5 6
Figura 11. Seções transversais da região nodal de Talinum paniculatum, mostrando em seqüência a
vascularização da folha (seta) e da gema axilar (asterisco e ponta de seta). Escala = 200µm.
1 2
1 2
2 1 1
3 4
Figura 12. Seções transversais da região nodal de Talinum paniculatum, mostrando a vascularização da
gema axilar e das estruturas formadas (setas, A). Notar que o sistema vascular das estruturas está
presente na base (C) e praticamente ausente em regiões mais apicais (D). 1 = par de profilos; 2 =
estrutura abaxial. Escala = 200µm (A, C), 100µm (B, D).
1
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1 2
Mm
En
Mm
3 4
Mm Mm
En
5 6
Figura 1. Seções transversais do ápice caulinar de Portulaca werdermannii e P. halimoides.

Primórdios foliares em diferentes estágios de desenvolvimento em P. werdermannii (A) e P.
halimoides (B). Detalhes de primórdios foliares, mostrando estágios de diferenciação do
meristema marginal (Mn) e da endoderme (En) em P. werdermannii (C, E-F) e P. halimoides
(D). Notar divisão periclinal da hipoderme (ponta de seta). Barras = 4007m (A), 2007m (B, F),
307m (C), 507m (D-E).
Hp
Hp
1 2
En
En
3 4
Vp
Figura 2. Seções transversais de primórdios foliares de Portulaca werdermannii e P.

halimoides. Estágios de diferenciação do sistema vascular periférico (Vp) e do fundamental
em P. werdermannii (D-E) e P. halimoides (A-C). Notar hipoderme já diferenciada em A e B
(Hp) e células da endoderme com divisão periclinal (En). Célula contendo drusa (seta branca)
Barras= 507m (A-D), 2007m (E).
1 2 En
Pa
3 4 En
5
Figura 3. Seções transversais da folha de Portulaca halimoides (A-B, E) e P. werdermannii
(C-D). Neste estágio, não se observam regiões meristemáticas. Notar o amplo parênquima
aquífero (Pa) e a anatomia Kranz (endoderme, En) associada ao mesofilo radiado (ponta de
seta). Células contendo mucilagem (seta). Barras= 4007m (A, C), 1007m (D-E), 507m (B).
1 2
En En
3 4
5 6
Figura 4. A-B. Seções transversais da folha de Portulaca halimoides (A) e P. werdermannii

(B). Detalhe do feixe vascular central sem a diferenciação da endoderme como bainha Kranz.
C-F. Seções transversais da região basal da folha de P. halimoides (C,D) e P. werdermannii
(E,F) mostrando a redução (em direção à base) das porções meristemáticas que darão origem a
novos feixes periféricos (região tracejada). Notar divisões periclinais das células da endoderme
(En). Barras= 507m (A, C-D), 1007m (B, E-F).
*
*
1 * 2
*
Mn
3 4
Mn
Mn
5 6
Figura 5. Seções transversais do ápice caulinar de Talinum paniculatum (A,C,D), Portulaca

mucronata (E) e P. hirsutissima (B,F). A-B. Primórdios foliares em diferentes estágios de
diferenciação e base de folhas (asterisco). C-D. Detalhe dos primórdios foliares de T.
paniculatum, sendo possível observar que as células da porção abaxial estão vacuoladas, em
contraste com as células da face adaxial que apresentam citoplasma denso (meristema
marginal, Mn). E-F. Primórdios foliares em estágios de diferenciação semelhante ao observado
na figura D. Barras= 200µm (A, B), 50µm (C,F),100µm (D, E).
En
En
1 2
3 4
5 6
Figura 6. Seções transversais de primórdios foliares de Talinum paniculatum (A), Portulaca

mucronata (B-D, F) e P. hirsutissima (E). Notar nas figuras A e B, endoderme com atividade
mitótica (En). Nas figuras B-F é possível observar que células derivadas do meristema
marginal se dividem em diferentes planos, resultando na formação de cordões procambiais
apenas na face adaxial, caracterizando uma simetria dorsiventral. Barras= 50µm (A-D, F),
100µm (E).
1 2
3 4
5 6
Figura 7. Seções transversais de primórdios foliares de Portulaca mucronata (A,D), Talinum

paniculatum (B,E) e P. hirsutissima (C,F). Notar diferentes atividades do meristema marginal:
em P. mucronata (A) há atividade uniforme em ambas as faces; T. paniculatum (B) com
intensa divisão celular na face adaxial e P. hirsutissima (C) com atividade do meristema
marginal acentuada na face adaxial, e reduzida no sentido lateral. D-F. Estágios posteriores de
diferenciação da lâmina foliar. Neste estágio, com o crescimento diferencial de ambas as faces
da lâmina foliar, as folhas de T. paniculatum se distendem e perdem a conformação revoluta
(E). Nos feixes laterais não foi observada anatomia Kranz (D-F). Células de mucilagem foram
observadas em toda região do mesofilo (setas). Barras= 200µm (A), 100µm (B, D), 50µm (C,
E-F).
*
1 2
Figura 8. Seções transversais da região da base da folha de Talinum paniculatum (A,

asterisco), Portulaca mucronata (B) e P. hirsutissima (C). Notar organização radiada do
sistema fundamental na região adaxial resultante de divisões periclinais (A-B). Barras= 100µm
(A-C).
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Desenvolvimento da folha e axila foliar em

Leaf and leaf axil development in Portulacaceae

Desenvolvimento da folha e axila foliar em

Leaf and leaf axil development in

Dissertação apresentada ao Instituto

Orientadora: Profa. Dra. Gladys

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D9223D '2TTS)D 325B5D D 276AD A24FAD 2D !55252D 2D 3BA7D

75D5472D4AD;2D5AD62DG56AD !KCDA356ADB25D3=AD 455B723A252CD

5D 5>5D !55252D 2D A7D ,423A7D 123456787F 2D 976AB5C"D D 5472D A=4565D 657D

6232427D 72;[,457D B5657D 7232D ;2CD 27AD ;2D 5B2457D A623565242D

772456ACD !55252D 622D 23D AAD 3BAD 3&AD 5D 3=AD 455B723A252CD

5QAD 62D 26,457D 5BA4546AD B535D 5D A46%&AD B535F5D 62D A355252CD

62D 5D 7B235>5D A4246AD !55252D 2D 3,7D 5>57D A4A24F357D ;2D

A3AB57AD ':"D 2)"D D272D 35=5AD A7D 5A327D 6272=35D GA355252KCD

5D 7=A362D A354252D FD ABA75D BA3D AAD 5>57D 455B723A5252D

'B787CD32!F 377CA7D 2D 976AB2A)CD +5725252CD !55252CD 62325252D

'446AD 764324D87CD D373A7D 2D 1234567873A7)CD 15AB95252CD (A45252D

'446AD 12A325252CD 767B3ABA7CD A47B4DCD 6742BA7CD #D$AA7CD %2B4A7CD

D =75D B25D 625%&AD 657D 325%H27D A24F57D 45D 7=A362D

A354252D 2545D A35D ;27&AD 423277542D AD 276AD 65D 2A%&AD 6A7D

B53,4;5D A2AD A4246AD A23345CD B53D 4234AD 62D =35FA57CD B2354AD

WD 23A7AD FD 5242D 32A426ACD 27B25242D 2D !55252D I57D

B232427D AD A57D 72457D B23772427CD A3546AD 3A7257D '#D$AA7)1D '2)D

23=257D B232427D AD 35>B27D 534A757D 2D A57D 326B657D AD 72457D

'B787CD32F 2D 2BA7)&F '3)D 23=257D 5457D AD 527D B3A7356A7D 2D A57D

72457D '123456787)1D '4)D B2;24A7D 53=7A7D AD 3A327D B5;5272427D AD

23=257D '976AB5C)1D '6)D 32B562357D BAAD 72457D 2D =572D 27B277565D '(7D667)D

WD 7324AD 62D 6232427D 2735F57D 565B557D 5AD 5=242D F3AD 2D

3BA7D &AD B3<A7D A2425242D 627B23AD AD 4232772D 62D 6237A7D

A57D AD 7225427D J;257D A242D 24A435657D 457D 6257D

26A26N4257CD 5B32724546AD =AD 2B5466AD 2D B2>AAD '+AX2CD 0#541D 8=7A4D VD

D 3547%&AD 2432D B32724%5D 62D A57D GA47KCD '1D3DA7)CD 2D 57,45D 62D

'WB4A6252D 2D%7A5DBA7)CD 2AD A7D B27;756A327D 5D A532D B535D 57D7=5>57D

!A46ACD 322427D 276A7D 24A246AD !55252D 2D 727D 3BA7D 57D

!A46ACD A3A7D 5352327D 4232775427D AA332D 45D 55D A53D 2D

6232427D 27BF27D 6AD 3BA"D !55252CD BA3D 22BACD 5B327245D 35A7D

=575CD 5565D 62D 23725D 423553D 'GAB4D387673F CD3A4DCK)D AD BA45D

WB4A6252"D @757D 273357D 7&AD 627357D AAD B2;24A7D 27B4A7D 2D A356A7D

+AX2D '0#44)D A=7235D 5D B32724%5D 62D <3&A7D 423263A7D 2432D 27B4A7D 2D

27B4272427D 5D <3&A7D B355242D AA7A7"D 7D A357D 57D 42326357D

27B4A7D 2365623A7D BA62D 5B53223D 427757D A357D 42326357D 25D

2D!55252CDBAA7D276A7D4275DA3A7D3BA7D6AD56AD !"D D55D62D

5A327D 4A35%H27CD 7A=326AD ;2D 5=A362D 5D 545A5D AD D 24A;2D

A24FACD A47623546AD 57D 322427D 654%57D 45D AB32247&AD 657D 325%H27D

4A7B23D9A249D83ABD' 8)"D2TT#"D 4DB652DAD2D 4A7B23D9A249D

!3A4;7CD "D 0#6$"D *CAC.5E7352A D26A EDD352A 5AE5/C42BA ED23"D !A=5D

D9223CD A"D VD @CD -"D 2T0T"D 742354D A355252D D 42PD 55D

A342CDA"D:"D0#S6"D DB9A242D5775A4DAD2D 4A7B2352"D0.5E7352D4A

D 5>5D A355252CD AAD 32A4265D 356A45242CD 55D 76AD

A427565D AAD A4AF5D 623757D 2B27D '!53A4D 0#$S1D 1237XABCD 0##31D

557D 6257D A35D D 56AD =2D 772456ACD 624A456AD !8

47D 5352327D A3A8545NA7D B3272427D 4272D 3BAD B5322D BA773D

!D ' 455B723A252D ZD !55252D ZD 123456787)CD 7246AD 5D 657D BA57D

AA7D 35=5A7CD 5FD 62D !A347X9D '0#30)CD 3255D 5D 27335D 62727D

3A57D 2D GA355252K"D @=A35D 275D 6232427D BA7CD AD 535%H27D

5B327245D 72B32D 5D 275D 27335D =75D 2D 72;[,45D 62D 62724A24ACD

7246AD 72B32D6273A7DAAD 772356A7D B535D AD 3BAD '35=5X53D VD A55995CD

ED B535D 5D5>5D !55252DD6237A7D35=5A7D ;2D 3255D 5DAA33,45D 2D

D9223D VD @D '2T0T)D 2545D 5D B<272D 62D ;2D 5D AA33,45D 62757D