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Artigo : Abordagem ecofisiológica dos manguezais-Uma revisão

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Prévia do material em texto

Revista Biotemas, 27 (3), setembro de 2014
1Biotemas, 27 (3): 1-11, setembro de 2014
ISSNe 2175-7925 
http://dx.doi.org/10.5007/2175-7925.2014v27n3p1
Revisão
$ERUGDJHP�HFR¿VLROyJLFD�GRV�PDQJXH]DLV��XPD�UHYLVmR
Sávia Soares Pascoalini 1
Dielle Meire de Santana Lopes 2
Antelmo Ralph Falqueto 2
Mônica Maria Pereira Tognella 2*
1 Universidade Federal do Espírito Santo, Vitória – ES, Brasil 
2 Universidade Federal do Espírito Santo, Centro Universitário Norte do Espírito Santo
Rodovia BR 101 Norte, Km 60, CEP 29932-540, São Mateus – ES, Brasil 
* Autor para correspondência
monica.tognella@gmail.com
Submetido em 30/11/2013
Aceito para publicação em 26/05/2014
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O manguezal apresenta elevada produtividade primária que é, em parte, resultado dos mecanismos 
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DUWH�GRV�HVWXGRV�HFR¿VLROyJLFRV�VREUH�RV�PDQJXH]DLV�H�LGHQWL¿FDU�ODFXQDV�TXH�SRVVLELOLWHP�DPSOLDU�R�FRQKHFLPHQWR�
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espécies. A resposta de determinada espécie ao estresse depende de sua tolerância. Conclui-se que os estudos 
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Key words:�)ORRGLQJ��1XWULHQWV��3KRWRV\QWKHVLV��6DOLQLW\
Revista Biotemas, 27 (3), setembro de 2014
2 S. S. Pascoalini et al.
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para os manguezais mundiais. Para isto, realizamos 
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Os manguezais despertam interesse de especialistas 
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elevada produtividade (POOP et al., 1985; BALL, 1988; 
NAIDOO et al., 2002; 2011; PARIDA et al., 2004; 
BARR et al., 2009).
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(VWXGRV� UHODFLRQDGRV� j� REWHQomR�GH� GDGRV� in situ de 
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climáticas.
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SRU�HVVDV�HVSpFLHV�YHJHWDLV�VmR�R�IRFR�GHVWD�UHYLVmR��QD�
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SDUD�DV�PXGDQoDV�FOLPiWLFDV��
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As espécies de mangue são consideradas plantas 
C3, pois não apresentam anatomia de plantas C4, sua 
FRPSRVLomR�LVRWySLFD��į13C) varia entre 24,6 e 32,2 %o, 
VHX�SRQWR�GH�FRPSHQVDomR�GR�&22�H�VXD�WD[D�Pi[LPD�
GH� IRWRVVtQWHVH� HP� WHPSHUDWXUD� DEDL[R� GH� ��ƒ&� VmR�
características consideradas típicas de plantas C3 (BALL; 
)$548+$5������D�� ����E��%$//�� ������3$5,'$�
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FRPSHQVDomR�GR�&22 característicos das C3.
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de trocas gasosas. Estas tratam da entrada de CO2 para 
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(BALL, 1988; NAIDOO et al., 2002).
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Avicennia germinans (L.) L., Laguncularia racemosa 
(L.) Gaertn. e Rhizophora mangle L., sendo que as 
SULPHLUDV�REWLYHUDP�PDLRU�DVVLPLODomR�GH�FDUERQR�GR�
que R. mangle menos tolerante ao sal. O autor relaciona 
R�PDLRU�GHVHPSHQKR�IRWRVVLQWpWLFR�UHDOL]DGR�SRU�DTXHODV�
HVSpFLHV�DR�PHOKRU�XVR�GH�iJXD�HP�QtYHO�IROLDU��DSHVDU�
de apresentarem menor transporte de água.
Revista Biotemas, 27 (3), setembro de 2014
3$ERUGDJHP�HFR¿VLROyJLFD�GRV�PDQJXH]DLV��XPD�UHYLVmR
6DOLQLGDGH��VHXV�HIHLWRV�VREUH�D�SODQWD
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deve-se, principalmente, ao cloreto de sódio (NaCl) 
dissolvido na água, correspondente a 86% do total. A 
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���J�NJ��FRUUHVSRQGHQGR�D�����PM de Na+ e 558 mM 
de Cl-��/,%(6���������e�FRQVLGHUDGR�R�SULQFLSDO�IDWRU�
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�3$5,'$��-+$���������'HVVD�IRUPD��GLYHUVRV�HVWXGRV�
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Estas espécies apresentam comportamento distinto 
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parviflora �5R[E���:W�� 	�$UQ�� H[� *ULII� tem seu 
Pi[LPR� FUHVFLPHQWR� HP� VDOLQLGDGH� HP� WRUQR� GH� ��
(PARIDA et al., 2004). Rhizophora mucronata Lam. 
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GH���D�����+233(�63((5�HW�DO���������HP�ODERUDWyULR��
Avicennia marina� �)RUVN���9LHUK��� FRQVLGHUDGD�PDLV�
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VDOLQLGDGH�GH��� �&/28*+��������63$/',1*�HW� DO���
2010).
Em geral, o mangue tem seu desenvolvimento 
yWLPR�HP�EDL[D�VDOLQLGDGH��&RQWXGR��DOJXPDV�HVSpFLHV�
toleram salinidade mais elevada do que outras (BALL, 
1988). Espécies do gênero Rhizophora são mais 
sensíveis ao aumento de salinidade do que aquelas do 
gênero Avicennia �&/28*+���������$V�GLIHUHQoDV�HVWmR�
presentes dentro dos gêneros. Rhizophora apiculata 
Blume é mais sensível do que Rhizophora stylosa�*ULII��
�%$//�HW�DO���������
É notório que as espécies de manguezal são 
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espécie para espécie dependendo da tolerância.
Normalmente, manguezais diminuem o potencial 
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DXPHQWD��RFRUUH�HIHLWR�VLPLODU�QDV�FRQFHQWUDo}HV�GH�1D+ 
e Cl-��(VWH�IDWR�QmR�p�DFRPSDQKDGR�SHOD�HQWUDGD�GH�tRQV�
K+�SDUD�HVSpFLHV�PHQRV�WROHUDQWHV�j�VDOLQLGDGH��(VWXGRV�
YHUL¿FDUDP�DXPHQWR�QD� UD]mR�1D+/K+ para Bruguiera 
gymnorrhiza (L.) Lam. crescendo em 35 de salinidade. 
Entretanto, Avicennia marina mantém menor potencial 
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zonas como proposto por Davis Jr. (1940).
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),/+2�� ������� ([SHULPHQWRV� UHDOL]DGRV� SRU� 3DULGD�
et al. (2003), com plântulas de Bruguiera parviÀora 
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VDOLQLGDGH� Ki� SHUGD� SDUFLDO� GR� HPSLOKDPHQWR� JUDQDO��
GR� FLWRFURPR� %�I� H� GLVVRFLDomR� GH� SROLSHSWtGLRV�
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GLVIXQo}HV�DIHWDP�GLUHWDPHQWH�D�DTXLVLomR�GH�FDUERQR�
H�� LQGLUHWDPHQWH�� R� LQFUHPHQWR� HP� ELRPDVVD��$OpP�
GLVVR�� KLSHUVDOLQLGDGH� SRGH� FDXVDU� GHVLQWHJUDomR� GH�
FORURSODVWRV�H�GH�PLWRF{QGULDV��FRPR�REVHUYDGR�HP�A. 
marina�VXEPHWLGD�D����GH�VDOLQLGDGH��1$,'22�HW�DO���
2011).
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D� FRQFHQWUDomR� GH� FORURILOD�� R� TXH� SRGH� UHGX]LU� D�
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1R� HQWDQWR�� 6REUDGR� ������� REVHUYRX� DXPHQWR� QD�
FRQFHQWUDomR� GH� FORURILOD� a e b em Laguncularia 
racemosa,�R�TXH�LQGLFD�WROHUkQFLD�jV�FRQGLo}HV�LPSRVWDV�
SHOD�FRQFHQWUDomR�GH�VDLV�TXH��QR�H[SHULPHQWR��QmR�IRUDP�
superiores a 30. Normalmente, o aumento da salinidade 
YHP� DFRPSDQKDGR� SHOD� UHGXomR� QD� FRQFHQWUDomR� GH�
FORUR¿OD�GHYLGR�j�IRWRLQLELomR�FU{QLFD��SRLV�R�WHRU�VDOLQR�
é superior ao tolerado para a espécie estudada, como 
YLVWR� QRV� WUDEDOKRV� GH�6REUDGR� ��������1DLGRR� HW� DO��
(2002; 2011) e Parida et al. (2003).
Revista Biotemas, 27 (3), setembro de 2014
4 S. S. Pascoalini et al.
7RGRV� HVWHV� IDWRUHV� DVVRFLDGRV� j� GHVDWLYDomR�GR�
)6,,� OHYDP� j� GLPLQXLomR� GD� DWLYLGDGH� IRWRTXtPLFD��
$� UHGXomR� GD� HILFLrQFLD� IRWRTXtPLFD� �)v/Fm) com 
LQFUHPHQWR�HP�VDOLQLGDGH�p�REVHUYDGD�SRU�YiULRV�DXWRUHV�
para A. marina, B.parviÀora e R. mucronata (PARIDA 
HW�DO���������+233(�63((5�HW�DO���������1$,'22�HW�
DO����������2�HVWUHVVH�VDOLQR�SURYRFD�WDPEpP�UHGXomR�
GD�FRQGXWkQFLD�HVWRPiWLFD�H�GD�FRQGXomR�QR�PHVR¿OR��
OLPLWDQGR� D� IRWRVVtQWHVH� SRU� EDL[D� GLIXVmR� GR�&22 
(PARIDA et al., 2004), além de diminuir condutividade 
KLGUiXOLFD� SURYRFDQGR� R� IHFKDPHQWR� GRV� HVW{PDWRV�
(SOBRADO, 2001). 
(VWUDWpJLDV� GH� DEVRUomR� GH� iJXD� HP�
FRQGLo}HV�VDOLQDV
0HVPR�FUHVFHQGR� HP�XP�DPELHQWH� LQXQGDGR�� D�
VDOLQLGDGH� HOHYDGD�GL¿FXOWD� D� DEVRUomR�GH� iJXD�SHODV�
SODQWDV��3$/,<$987+�HW�DO����������$VVLP��DV�HVSpFLHV�
GH�PDQJXH� WrP�PHFDQLVPRV� SDUD� DEVRUYHU� D� iJXD�
FRQWUD� IRUWHV� SRWHQFLDLV� RVPyWLFRV� �720/,1621��
�������&UHVFHU� HP�XP�PHLR� VDOLQR� FULD� EDVLFDPHQWH�
GRLV�SUREOHPDV�SDUD�D�SODQWD������D�UHJXODomR�RVPyWLFD�
H�����D�WR[LGDGH�GHVWHV�tRQV��$VVLP��DV�SODQWDV�WrP�GH�
DEVRUYHU�RV�tRQV�LQRUJkQLFRV�SDUD�QHXWUDOL]DU�R�JUDGLHQWH�
RVPyWLFR��7RGDYLD�� R� H[FHVVR� GHVWHV� QR� LQWHULRU� GD�
FpOXOD� WRUQD�VH� Wy[LFR�H� WHP�HIHLWRV�QHJDWLYRV�VREUH�D�
UHSURGXomR��R�FUHVFLPHQWR�H�VREUHYLYrQFLD�GDV�SODQWDV�
�0(+/,*��������
3DUD�PDQWHU� D� DEVRUomR� GH� iJXD�� DV� SODQWDV� GH�
mangue não apenas precisam restringir a perda de água 
SRU�PHLR� GH� DGDSWDo}HV�PRUIROyJLFDV� H� ¿VLROyJLFDV�
�HVSHVVDPHQWR� IROLDU�� DXPHQWR� GR� WHPSR�GH� UHWHQomR�
GR�QLWURJrQLR�H�H¿FLrQFLD�QR�XVR�GD�iJXD���PDV�WDPEpP�
PDQWHU� RV� SRWHQFLDLV� GH� iJXD� VX¿FLHQWHPHQWH� EDL[RV�
(LOVELOCK; FELLER, 2003; KRAUSS et al., 2008) 
QR� LQWHULRU�GDV� IROKDV��(VWHV�DVSHFWRV� ID]HP�FRP�TXH�
D�H¿FLrQFLD�GH�XVR�GD�iJXD��HP�DOJXPDV�GDV�HVSpFLHV�
GH�PDQJXH�� VHMD� FRQVLGHUDGD� XPD� GDV�PDLV� DOWDV� GH�
todas as plantas C3 (BALL, 1986), uma vez que estas 
QRUPDOPHQWH�DSUHVHQWDP�EDL[DV�WD[DV�GH�WUDQVSLUDomR�H�
FRQGXWkQFLD�HVWRPiWLFD��%$//��)$548+$5������D��
����E�� %$//�� ������ /29(/2&.�� %$//�� ������
SOBRADO, 2005).
Entretanto, apesar dessas estratégias, para 
compensar a elevada osmolaridade de sal no solo, um 
GRV�PHFDQLVPRV�ELRTXtPLFRV�HPSUHJDGRV�p�R�DF~PXOR�
de solutos compatíveis, a partir do aumento de moléculas 
SHTXHQDV�GH�RVPyOLWRV�FRPR�D�JOLFLQD�EHWDtQD�RX�iOFRRLV�
GH�Do~FDU��3233�HW�DO���������6$.$0272��085$7$��
2000) que ampliam o potencial osmótico sem alterar o 
L{QLFR��5HGXo}HV�QRV�SRWHQFLDLV�RVPyWLFRV�DWUDHP�iJXD�
para dentro da célula e permitem manter o potencial 
GH�SUHVVmR��02*+$,(%�HW�DO���������QHFHVViULR�SDUD�
REWHU�iJXD�
$�PHOKRU�H¿FLrQFLD�GR�XVR�GD�iJXD�DVVRFLDGD�DRV�
FXVWRV� HQHUJpWLFRV� GH� H[FUHWDU� H�RX� H[FOXLU� RV� VDLV� p�
j�FXVWD�GH� WD[DV� UHGX]LGDV�GH�DVVLPLODomR�GH�FDUERQR�
(BALL, 1988). Quanto maior o teor de sal no local 
PHQRU�VHUi�D�SURGXWLYLGDGH�H�LVWR�LUi�UHÀHWLU�QD�HVWUXWXUD�
GR�ERVTXH�
Durante os períodos de elevada demanda 
HYDSRUDWLYD�� SODQWDV� TXH� H[FOXHP� VDO� SUHFLVDP�
economizar água por causa da capacidade limitada para 
H[WUDt�OD�GRV�VRORV�VDOLQRV��%$55�HW�DO����������6PLWK�HW�
DO���������PRVWUDUDP�TXH�D�UD]mR�SDUD�D�UHGXomR�GD�WD[D�
GH�DVVLPLODomR�GH�FDUERQR�GR�PDQJXH�QD�9HQH]XHOD�IRL�
HP�SDUWH�GHYLGR�j�VDOLQLGDGH�HOHYDGD��R�TXH�FDXVRX�PDLRU�
SRWHQFLDO�RVPyWLFR�QR�[LOHPD�H�UHGXomR�GD�FRQGXWkQFLD�
HVWRPiWLFD��(P�DPELHQWHV�KLSHUVDOLQRV��RV�PDQJXH]DLV�
DSUHVHQWDP�GLPLQXWDV�WD[DV�GH�DVVLPLODomR�GH�FDUERQR�
H�FRQGXWkQFLD�HVWRPiWLFD��RULJLQDQGR�ERVTXHV�GHVFULWRV�
FRPR�DQ}HV��/8*2��61('$.(5��������
2XWUR� SUREOHPD� HQIUHQWDGR� SHORV�PDQJXH]DLV��
GHYLGR�j�VDOLQLGDGH�H�D�HOHYDGD�GHPDQGD�HYDSRUDWLYD��
p�VXD�YXOQHUDELOLGDGH�j�HPEROLD�RX�FDYLWDomR�TXH�SRGH�
REVWUXLU�D�FRQGXWkQFLD�KLGUiXOLFD��(VVD�YXOQHUDELOLGDGH�p��SULQFLSDOPHQWH��SUHVHQWH�HP�DPELHQWHV�KLSHUVDOLQRV��
SRLV� DSUHVHQWDP� GLILFXOGDGH� HP� DEVRUomR� GH� iJXD�
�/29(//2&.�HW�DO�������D������E��0$57,1�HW�DO���
������� (VWXGRV� GH� 6REUDGR� �������� FRP�Avicennia 
germinans, demonstram que as espécies com maior 
H¿FLrQFLD� QR� XVR� GD� iJXD� SRVVXHP� HVWD� FDSDFLGDGH�
GHYLGR�j�UHGXomR�QD�FRQGXomR�GH�VHLYD�QR�FDXOH��(VWD�
estratégia é possível porque as espécies reduzem a 
FRQGXWkQFLD�HVWRPiWLFD�D�SRQWR�GH�LPSHGLU�D�HPEROLD�
QDV�FpOXODV�GR�[LOHPD�
Revista Biotemas, 27 (3), setembro de 2014
5$ERUGDJHP�HFR¿VLROyJLFD�GRV�PDQJXH]DLV��XPD�UHYLVmR
7RPOLQVRQ���������DR�GHVFUHYHU�R�WHFLGR�[LOHPiWLFR�
GR�PDQJXH��Mi�DVVRFLD�FRPR�PHFDQLVPRV�SDUD�HYLWDU�D�
HPEROLD�D�GLPLQXLomR�QR�GLkPHWUR�FHOXODU��+DR�HW� DO��
�������� FRPSDUDQGR� D� SUHVVmR� RVPyWLFD� QR� VROR� VRE�
GLIHUHQWHV�VDOLQLGDGHV�HP�SHUtRGRV�GLXUQRV�H�QRWXUQRV�
para Rhizophora mangle��LGHQWL¿FDP�TXH�D�UHGLVWULEXLomR�
GD�VHLYD�EUXWD�RX�UHYHUVmR�GD�FRQGXomR�RFRUUH�SDUD�HYLWDU�
D�HPEROLD��DOpP�GH�DPHQL]DU�D�VDOLQLGDGH�QR�VHGLPHQWR�
HP�VLWXDo}HV�GH�EDL[D�IUHTXrQFLD�GH�LQXQGDomR��
0HFDQLVPRV�GH�WROHUkQFLD�j�VDOLQLGDGH
$V�HVWUDWpJLDV�SDUD�GHWHUPLQDU�EDL[DV�WD[DV�GH�SHUGD�
GH� iJXD� IROLDU� WDPEpP� OLPLWDP�R� JDQKR�GH� FDUERQR��
ID]HQGR�FRP�TXH�HVSpFLHV�TXH�WrP�DPSOD�WROHUkQFLD�DR�
VDO�FUHVoDP�PDLV�OHQWDPHQWH�FRP�DXPHQWR�GD�VDOLQLGDGH�
do que aquelas menos tolerantes (BALL, 1986; 1988).
De acordo com suas características, as espécies de 
PDQJXH�SRGHP�VHU�FODVVL¿FDGDV�HP�VHFUHWRUDV�GH�VDO�±�
aquelas que têm glândulas de sal ou pelos para remover o 
sal – e não secretoras – que não apresentam mecanismos 
SDUD�D�H[FUHomR�GR�VDO� �6&+2/$1'(5�HW�DO���������
720/,1621�� ������� /DUFKHU� ������� GHVFUHYH� DV�
KDOy¿WDV�FRPR�SODQWDV�TXH�WROHUDP�RX�TXH�UHVLVWHP�DR�
sal. Aquelas que resistem ao sal apresentam mecanismos 
¿VLROyJLFRV�TXH�VmR��HYLWDU�D�HQWUDGD��GLOXLU�H�HOLPLQDU�R�
VDO��$V�FODVVL¿FDGDV�FRPR�WROHUDQWHV�VHULDP�DTXHODV�TXH�
WROHUDP�R�VDO�QR�FLWRSODVPD��'H�DFRUGR�FRP�6DOLVEXU\�
H�5RVV���������HVWDV�VHULDP�DV�KDOy¿WDV�YHUGDGHLUDV��RX�
HXKDOy¿WDV���6RPHQWH�DV� WROHUDQWHV�VHULDP�FDSD]HV�GH�
WROHUDU�RV�HIHLWRV�Wy[LFRV�GR�FORUR�H�VyGLR�QR�FLWRSODVPD�
�/$5&+(5��������6$/,6%85<��5266��������
Popp et al. (1993), revisando os conceitos propostos 
SRU� 6FKRODQGHU� ������� H� D� FODVVL¿FDomR� GH�/DUFKHU�
��������DYDOLDP�D�FDSDFLGDGH�GD�YHJHWDomR�GH�PDQJXH�
como capaz de evitar a entrada de sais via raiz e as 
HVWUDWpJLDV�GH�H[WUXVmR�H�RX�FRPSDUWLPHQWDomR�GRV�VDLV��
&RQFOXHP�TXH�RV�HQWHQGLPHQWRV�VREUH�RV�SURFHVVRV�GH�
H[FOXVmR�GR�VyGLR�H�FORUR�DQWHV�GR�LQJUHVVR�QR�[LOHPD�
QmR�VmR� WRWDOPHQWH�HQWHQGLGRV��KDYHQGR�HVSpFLHV�TXH�
QmR�GHPRQVWUDP�GLIHUHQoDV�HQWUH�DV�SUHVV}HV�RVPyWLFDV�
GR�VHGLPHQWR��GDV�UDt]HV��GRV�FDXOHV�H�GDV�IROKDV�FRPR�
D¿UPDGR�SRU�6FKRODQGHU���������3RU�RXWUR�ODGR��D¿UPDP�
TXH� DV� HVSpFLHV�TXH�SURPRYHP�D� H[WUXVmR�GR� VDO� YLD�
glândulas são reduzidas dentro da gama de espécies de 
mangue.
(P� UHODomR� j� FRPSDUWLPHQWDomR� GRV� VDLV�� HVWH�
IHQ{PHQR�p�GHVFULWR�VHQGR�FRPXP�HQWUH�DV�HVSpFLHV�GH�
PDQJXH�� Mi�TXH�YiULDV�DSUHVHQWDP�PDLRU�FRQWH~GR�GH�
VDLV�QDV�IROKDV�VHQHVFHQWHV�TXDQGR�FRPSDUDGDV�jV�QRYDV�
(POPP et al., 1993). Além disso, estudos realizados pelos 
autores revelaram que com o aumento da salinidade 
RFRUUH�DFUpVFLPR�GH�PHWDEyOLWRV�VHFXQGiULRV��FRPR�R�
manitol, no citoplasma, concomitante ao incremento de 
sódio e cloro nos vacúolos.
Os gêneros Aegiceras, Avicennia, Acanthus e 
Aegialitis WrP� HVWUXWXUDV� JODQGXODUHV� GH� H[FUHomR� GR�
sal enquanto Laguncularia e Conocarpus têm estrutura 
DQiORJD� jV� JOkQGXODV� GH� VDO� �720/,1621�� �������
Bruguiera gymnorrhiza e Rhizophora mucronata 
WDPEpP�H[FOXHP�VDO��QR�HQWDQWR��QXP�SURFHVVR�GH¿QLGR�
FRPR�UHDORFDomR�GRV�VDLV�SDUD�SDUWHV�VHQHVFHQWHV��RQGH�
R�VDO�p�DFXPXODGR�QDV�IROKDV�PDLV�PDGXUDV��DV�TXDLV�VmR�
HOLPLQDGDV�SDUD�H[FOXLU�R�H[FHVVR� �67(,1.(���������
'LIHUHQWHV� HVSpFLHV� WrP�GLVWLQWDV� IDL[DV� GH� WROHUkQFLD�
ao sal (SUAREZ; MEDINA, 2005) correlacionadas a 
VXD�FDSDFLGDGH�GH�H[FUHomR�H�RX�HOLPLQDomR�GRV�VDLV��
$TXHODV� TXH� H[FUHWDP� RV� VDLV� GLDULDPHQWH� SRVVXHP�
FDSDFLGDGH� GH� FRORQL]DU� DPELHQWHV� FRP� VDOLQLGDGHV�
mais elevadas, ao contrário daquelas que armazenam 
RV� VDLV� SRU� SHUtRGRV�PDLRUHV� �SRU� FRPSDUWLPHQWDomR�
RX� UHDORFDomR�� Vy� HOLPLQDQGR�RV� FRP�PRUWH� GH� VXDV�
estruturas vegetais.
Avicennia�VmR�FRQKHFLGDV�SRU�H[FOXtUHP�FHUFD�GH�
����GH�VDO�DEVRUYLGRV�D�SDUWLU�GD�VXSHUItFLH�GD�UDL]��TXH�
SRGH�DXPHQWDU�j�PHGLGD�TXH�D�FRQFHQWUDomR�GH�VDO�GD�
iJXD�VH�DPSOLD��%5$1&+��%5$1&+���������$�H[FOXVmR�
GH�VDO�p�REWLGD�FULDQGR�VH�SUHVVmR�KLGURVWiWLFD�QHJDWLYD�
QD�FpOXOD�TXH�IRUQHFH�SUHVVmR�RVPyWLFD�VX¿FLHQWHPHQWH�
QHJDWLYD�SDUD�DV�UDt]HV�SDUD�DEVRUomR�GH�iJXD��HQTXDQWR�
WRGRV� RV� RXWURV� tRQV� VmR� H[FOXtGRV� �0221� HW� DO���
1986). Estes autores avaliaram que, além do processo 
GH�XOWUD¿OWUDomR��D�H[WUXVmR�GR�VDO�YLD�VLVWHPD�GH�UDt]HV�
para as espécies de Avicennia é controlada por aspectos 
DQDW{PLFRV�TXH� LQYLDELOL]DP�R� WUDQVSRUWH� DSRSOiVWLFR�
GD�iJXD�DWp�R�[LOHPD��(VWH�WUDQVSRUWH�p�LPSHGLGR�SHOD�
VXEHUL]DomR� GDV� FpOXODV� GR� FyUWH[� GD� UDL]� H� WDPEpP�
SHOD�EDQGD�GH�&DVSDU\�GD�HQGRGHUPH��6XDUH]�H�0HGLQD�
Revista Biotemas, 27 (3), setembro de 2014
6 S. S. Pascoalini et al.
(2005) mostraram que plântulas de A. geminans�IRUDP�
VLJQL¿FDWLYDPHQWH� DIHWDGDV� SHOD� VDOLQLGDGH� H� WLYHUDP�
WROHUkQFLD� DR� VDO� HOHYDGD� HP�FRPSDUDomR� FRP�RXWUDV�
HVSpFLHV� VHPHOKDQWHV�� FRPR�Aegiceras corniculatum 
(L.) Blanco e Avicennia marina.
Estudo com Bruguiera gymnorhiza indicou que 
iUHD��PDVVD�IROLDU�SRU�SODQWD�H�ORQJHYLGDGH�DXPHQWDUDP�
FRP�R�LQFUHPHQWR�GD�VDOLQLGDGH��:$1*��/,1���������
$� VXFXOrQFLD� IROLDU� SRGH� VHU� XP�PHFDQLVPR� SDUD�
HYLWDU� FRQFHQWUDo}HV� GH� VDO� H[FHVVLYDPHQWH� HOHYDGDV�
�+$*(0(<(5�� ������� XPD� YH]� TXH� R� DXPHQWR� GR�
YROXPH�GR�YDF~ROR�SHUPLWH�D�GLOXLomR�GRV�tRQV�VRO~YHLV��
5HGXomR�GR�FUHVFLPHQWR��PHQRU�DOWXUD��GLPLQXLomR�
GD�ELRPDVVD�H�SURGXomR�GH�IROKD��SRGH�VHU�LQWHUSUHWDGD�
como resultado do investimento em mecanismos de 
tolerância e/ou resistência ao sal (SNEDAKER, 1982; 
68$5(=��0(',1$��������� Mi�TXH�D�PDQXWHQomR�GH�
tais estratégias reduz a produtividade primária líquida.
6HJXQGR� .DWKLUHVDQ� H� 7KDQJDP� �������� DV�
ÀXWXDo}HV�QD� VDOLQLGDGH�GHQWUR�GR� DPELHQWH�SDUHFHP�
WHU�HIHLWRV�PDLV�SURQXQFLDGRV�GR�TXH�D�KLSHUVDOLQLGDGH�
FRQWtQXD�� XPD� YH]� TXH� DV� SODQWDV� DGXOWDV� WrP� IDL[D�
VXSHULRU� GH� WROHUkQFLD� HP� UHODomR� jV� SOkQWXODV�� ,VVR�
DFRQWHFH�SRUTXH�DV�SODQWDV�PDLV�MRYHQV�VmR�JHUDOPHQWH�
PDLV� VHQVtYHLV� DR� VDO�� FUHVFHQGR�PHOKRU� HP� ORFDLV�
onde as salinidades são menores e, porteriormente, 
WRUQDP�VH�PDLV� WROHUDQWH� �6&+0,7=� HW� DO��� �������
(QWUHWDQWR��XPD�YH]�R�ERVTXH�HVWDEHOHFLGR��PRGL¿FDo}HV�
na salinidade podem levar ao colapso do indivíduo 
adulto por menor plasticidade que as plântulas, como 
REVHUYDGR� HP� VLWXDo}HV� RQGH� KRXYH� DOWHUDo}HV� QD�
GLQkPLFD� GH� LQXQGDomR�SURYRFDGDV� SRU�PRGL¿FDo}HV�
QD�GHVHPERFDGXUD�GR� ULR� �72*1(//$�HW�DO����������
2V�DXWRUHV�REVHUYDUDP�TXH�DV�DOWHUDo}HV�QD�VDOLQLGDGH�
WLYHUDP�FRPR�UHVSRVWD�GD�YHJHWDomR�D�PRUWH�GDV�IROKDV�
QD�SRUomR�PDLV�DOWD�GD�FRSD��5HODFLRQDUDP�WDO�IDWR�jV�
DOWHUDo}HV�GR�SRWHQFLDO�RVPyWLFR�TXH�SRGHP�WHU�RFRUULGR�
QRV�UDPRV�WHUPLQDLV�LQYLDELOL]DQGR�D�PDQXWHQomR�GDV�
IROKDV� SRU� GL¿FXOGDGH� QR� DSRUWH� GH� iJXD��&RP� LVWR��
os indivíduos adultos tiveram sua estatura diminuída, 
SHUPLWLQGR�VXD�VREUHYLYrQFLD��
,QXQGDomR�H�DODJDPHQWR
$V�ÀRUHVWDV�GH�PDQJXH]DO�VmR�UHJXODUPHQWH�ODYDGDV�
SHODV�PDUpV��$�FLUFXODomR�GD�iJXD�HOLPLQD�VXEVWkQFLDV�
Wy[LFDV�SDUD�D�YHJHWDomR��FRPR�R�VXOIHWR�GH�KLGURJrQLR�
�+26��H�R[LJHQDP�R�VHGLPHQWR��.1,*+7�HW�DO����������
2V�SDGU}HV�GH�LQXQGDomR�GDV�ÀRUHVWDV�VmR�GLIHUHQFLDGRV�
H�UHÀHWHP�QD�HVWUXWXUD�GD�PHVPD��2V�ERVTXHV�GH�IUDQMD�
são constantemente inundados pelo movimento da 
PDUp� YHQWLODQGR� H� UHPRYHQGR� RV�FRPSRVWRV� Wy[LFRV�
H� RV� ERVTXHV� GH� EDFLD� VmR�JHUDOPHQWH� LQXQGDGRV�QDV�
PDUpV�PDLV�DOWDV��6&+$())(5�129(//,�HW�DO���������
.1,*+7� HW� DO��� ������ ¿FDQGR�PDWpULD� RUJkQLFD� HP�
GHFRPSRVLomR�SRU�SHUtRGR�PDLRU��FDXVDQGR��HP�DOJXPDV�
VLWXDo}HV��FRQGLo}HV�GH�DQR[LD�SDUD�DV�UDt]HV�
(P�FRQGLo}HV�GH�DODJDPHQWR��R�PDQJXH�GHYH�OLGDU�
FRP�SDUFLDO�RX�FRPSOHWD�DXVrQFLD�GH�R[LJrQLR�GLVVROYLGR�
SDUD�UHVSLUDomR�GDV�UDt]HV��0&.((��������6.(/721��
$//$:$<���������(VVDV�HVSpFLHV��TXH�FDUDFWHUL]DP�D�
YHJHWDomR�GH�PDQJXH��GLIHUHP�HP�VXD�FDSDFLGDGH�GH�
PDQWHU�R�WHFLGR�GD�UDL]�DHUDGR�DIHWDQGR�D�¿VLRORJLD�H�
PRUIRORJLD�GDV�UDt]HV��0&.((���������$�KLSR[LD�FDXVD�
GLPLQXLomR�QD� WD[D�GH�H[WHQVmR�GD�UDL]�HP�Avicennia 
germinans e Laguncularia racemosa, mas não em 
Rhizophora mangle��$OpP�GLVVR��D�KLSR[LD�p�UHVSRQViYHO�
pela morte de raízes de L. racemosa H�SHOD�GLPLQXLomR�QR�
número de raízes laterais nas três espécies neotropicais 
FLWDGDV�DFLPD�H��WDPEpP��SHOR�DXPHQWR�GD�SDUWH�DpUHD�
em A. germinans e L. racemosa (MCKEE, 1996). 
Bruguiera mucronata apresentou similar tendência, com 
PDLRU�DOWXUD�TXDQGR�VXEPHWLGD�j�LQXQGDomR�FRQVWDQWH��
UHYHODQGR�PXGDQoD�QD�DORFDomR�GH�ELRPDVVD�SDUD�D�SDUWH�
DpUHD��+233(�63((5�HW�DO���������
Estudo realizado com plântulas de Avicennia marina 
DSRQWRX�TXH�R�HVWUHVVH�FDXVDGR�DSHQDV�SRU�LQXQGDomR�
SRGH� QmR� VHU� WmR� SUHMXGLFLDO� j� HVSpFLH� �6.(/721��
$//$:$<�� �������$V� SOkQWXODV� GH�A. marina em 
FRQGLo}HV� GH�PDUp� EDL[D� DXPHQWDP� D� FRQFHQWUDomR�
GH� R[LJrQLR� HP� VXDV� UDt]HV� DWp� FRQFHQWUDomR� HVWiYHO��
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3RUpP��D�FRQFHQWUDomR�UHVWDQWH�GH�R[LJrQLR�p�VX¿FLHQWH�
Revista Biotemas, 27 (3), setembro de 2014
7$ERUGDJHP�HFR¿VLROyJLFD�GRV�PDQJXH]DLV��XPD�UHYLVmR
SDUD�PDQWHU�PHWDEROLVPR� DHUyELFR� QR� QtYHO� GH� UDL]�
�6.(/721��$//$:$<�� ������� (VVD� FRQFHQWUDomR�
é determinada pela capacidade que as raízes têm de 
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DR�DHUrQTXLPD�QDV�UDt]HV��SRLV��HP�PDUp�DOWD��R�HVSDoR�
LQWHUQR�GH�JiV�HVWi�LVRODGR�GR�H[WHULRU�H�Ki�WUDQVIHUrQFLD�
GH�R[LJrQLR�GD�HVWUXWXUD�DpUHD�jV�UDt]HV��720/,1621��
������6.(/721��$//$:$<��������
(VWXGRV�HP�ODERUDWyULR�FRP�SOkQWXODV�PRVWUDUDP�
TXH� DV� LQXQGDo}HV� HP� ORQJR� SUD]R� QmR� WrP� HIHLWR�
SHUVLVWHQWH� VREUH� DV� WURFDV� JDVRVDV� HP�Avicennia 
germinans, Laguncularia racemosa ou Rhizophora 
mangle��3(=(6+.,�HW�DO���������.5$866�HW�DO����������
-i�D�LQXQGDomR�GH�FXUWR�SUD]R�UHGX]�D�DVVLPLODomR�Pi[LPD�
H�UHVSLUDomR�QR�HVFXUR�HP�SOkQWXODV�GH�A. germinans, 
L. racemosa e R. mangle de acordo com os estudos de 
Krauss et al. (2006). No entanto, os mangues maduros 
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do que as plântulas (NAIDOO et al., 1998). Apesar 
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UHVXOWDUHP�HP�WD[DV�GH�FUHVFLPHQWR�HOHYDGDV��HVWDV�VmR�
PHQRUHV�GR�TXH�DV�UHGXo}HV�QDV�WD[DV�GH�FUHVFLPHQWR�
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LQXQGDGRV��)$516:257+�HW�DO���������
1XWULHQWHV
O input de nutrientes no ecossistema manguezal é 
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GH� IUDQMD� VmR� H[SRVWRV� D� GXDV� FRQGLo}HV� QXWULFLRQDLV�
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do escoamento terrestre SDUD�PHOKRU�GHVHQYROYLPHQWR��
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Vy�RFRUUHQGR�QDV�PDUpV�PDLV�H[WUHPDV�GR�DQR��$OpP�
disso, apresentam salinidade acima daquela da água do 
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SURFHVVRV�PLFURELDQRV��FRPR�D�WD[D�GH�GHFRPSRVLomR�
H� UHWHQomR� GH� QXWULHQWHV� QD� UL]RVIHUD�� SRGHP� UHGX]LU�
D� GLVSRQLELOLGDGH�GH�QXWULHQWHV� SDUD� D� DEVRUomR�SHODV�
plantas (NAIDOO, 2006).
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KLGUiXOLFD�� GLPLQXLomR� GD� UDPL¿FDomR� GD� SDUWH� DpUHD�
e alongamento celular, reduzindo o crescimento das 
árvores. Portanto, em locais limitados por P os processos 
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HW�DO�������������D������E��1$,'22��������
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em altura das árvores. Esse elemento é utilizado na 
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SURGXomR� GH� VROXWRV� FRPSDWtYHLV� H� SDUWLFLSDP� QD�
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Revista Biotemas, 27 (3), setembro de 2014
8 S. S. Pascoalini et al.
(P�H[SHULPHQWRV�GH�ORQJR�SUD]R��GHPRQVWUDGR�SRU�
)HOOHU� �������HP�FRPXQLGDGHV�FUHVFHQGR� VREUH� VRORV�
FDOFiULRV�HP�7ZLQ�&D\V�%HOL]H��D�GLVSRQLELOLGDGH�GH�3�
IRL�R�IDWRU�HFROyJLFR�GHWHUPLQDQWH�SDUD�R�FUHVFLPHQWR�
GDV�SODQWDV��0HGLQD�HW�DO�� ������� WDPEpP�DVVRFLDUDP�
R� FUHVFLPHQWR� GH¿FLHQWH� GH�R. mangle, no leste de 
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H�HVWUHVVH�KtGULFR�VD]RQDO��/RYHORFN�HW�DO�������F���DR�
estudar manguezais onde P era o nutriente limitante, a 
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Estudando espécies de Avicennia gernimans 
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contrastantes na Australia, Martin et al. (2010) não 
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H� )HOOHU� �������� DR� DYDOLDUHP� D� IHUWLOL]DomR� GH�A. 
germinans versus Laguncularia racemosa constataram 
TXH� R� DXPHQWR� GR� GHVHPSHQKR� IRWRVVLQWpWLFR� HP�A. 
germinans com N era muito maior do que o de L. 
racemosa�IHUWLOL]DGR�FRP�R�PHVPR�QXWULHQWH�
2�QXWULHQWH�)RVIDWR� �3243-) em solos de mangue 
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limitados (REEF et al., 2010).
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Os ecossistemas costeiros, como o manguezal, 
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dos níveis de CO2, temperatura, nível médio do mar 
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aspectos possíveis que podem causar um impacto 
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FRPR�REVHUYDGR�HP�SOkQWXODV�GH�Rhizophora apiculata 
e Rhizophora stylosa FXOWLYDGDV�HP�ODERUDWyULR��%$//�
HW� DO��� �������1R� HQWDQWR�� GLIHUHQoDV� LQWHUHVSHFt¿FDV�
em resposta ao aumento de CO2�H�VXD�UHODomR�jV�RXWUDV�
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dentro de estuário (ALONGI, 2008).
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em áreas áridas e aumentaria em áreas de elevada 
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indivíduos de Rhizophora mangle�QD�)OyULGD��YHUL¿FDUDP�
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HW� DO�� ������� VXJHUHP� TXH� FHUFD� GH� ��� D� ���� GRV�
manguezais mundiais podem ser perdidos no período 
de 2000 a 2080, devido ao aumento em torno de 36 a 
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ressaltado por Alongi (2008) e Soares (2009). Além 
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Revista Biotemas, 27 (3), setembro de 2014
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as espécies (ALONGI, 2008).
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podem induzir respostas positivas do manguezal. Porém, 
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nível do mar e ocorrendo em velocidade acentuada, as 
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SODQWDV�GH�PDULVPDV�TXH� WHP�¿VLRORJLD�&4 e com isto 
assimilam CO2�GH�IRUPD�PDLV�H¿FLHQWH�HP�WHPSHUDWXUDV�
acima de 40ºC (ADAM, 1990).
Concluindo, esta análise reporta que os estudos 
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e, em alguns, apresentam resultados distintos entre 
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do manguezal ainda são incipientes sendo urgente 
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Os autores agradecem o apoio do Instituto Estadual 
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síntese. A primeira autora agradece a Pró-Reitoria de 
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)HGHUDO�GR�(VStULWR�6DQWR��SHORV�GRLV�DQRV�GH�EROVD�GH�
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de mestrado. A segunda autora agradece a CAPES pela 
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