I NFORMASJONSKILDER

Os peroxissomos são pequenas organelas, circundadas por uma membrana simples, envolvidas em reações metabólicas oxidativas (KAUR et al., 2009). Nos vegetais, há diversos tipos de peroxissomos que são especializados em certas funções metabólicas. Como exemplo os glioxissomos, peroxissomos foliares, peroxissomos de nódulos de raiz. Entretanto as diferenças entre os tipos de peroxissomos não são muito grandes, por isso, para simplificar, ficou definida a utilização do termo geral peroxissomos para todos os tipos de peroxissomos especializados (PRACHAROENWATTANA; SMITH, 2008)

Em 1966, De Duve e Baudhuin trabalhando com células de fígado de rato encontraram atividades de produção e degradação de H2O2 nessas organelas. Esta descoberta deu origem ao nome peroxissomos. Inicialmente acreditava-se que a função dos peroxissomos era apenas a remoção de H2O2, gerado na via fotorrespiratória,pela catalase. Entretanto diversos estudos atuais mostram que os peroxissomos participam de diversas funções, muitas delas envolvendo mais de uma organela. Como exemplo dessas funções o metabolismo dos lipídeos ( - oxidação), a síntese de hormônios vegetais, o ciclo do glioxilato, o metabolismo de ureídeos e os processos de desintoxicação celular (HAYASHI; NISHIMURA, 2003; KAUR et al., 2009).

Nos peroxissomos há produção de duas ERO, O2•− e H2O2, contudo, essa organela também possui um eficiente sistema de defesa antioxidante (Del RÍO et al. 2002, 2006) (Figure 8). Uma fonte de O2•− nos peroxissomos é a enzima xantina oxidase (XOD), uma xantina oxidoredutase oxidase que catalisa a oxidação da xantina ou hipoxantina a ácido uréico (SANDALIO et al., 1988; DEL RÍO et al, 1989; WERNER; WITTE, 2011). Outra via de produção de O2•− nos peroxissomos é através de uma pequena cadeia de transporte de elétrons na membrana do peroxissomo (STRUGLICS et al., 1993). O O2•− produzido tem meia vida de 2-4ms, antes de ser desprotonado a H2O2, espontaneamente ou através da enzima dismutase do superóxido (SOD), (VRANOVA et al., 2002). A SOD constitui a primeira linha de defesa, através da conversão de O2•− a H2O2, que pode ser metabolizado pela maquinaria antioxidante celular (NYATHI; BAKER, 2006).

Figura 8: Mecanismos de eliminação de ERO nos peroxissomos (Kaur et al. 2009).

Quanto ao H2O2, a glicolato oxidase é uma das principais enzimas responsáveis pela sua produção no processo de fotorrespiração (VOSS et al., 2012).A taxa de produção de H2O2 nos peroxissomos durante a fotorrespiração é duas vezes maior do que nos cloroplastos e cerca de 50 vezes maior do que na mitocôndria (FOYER; NOCTOR, 2003). A oxidação inicial de ácidos graxos através da enzima acyl-CoA oxidase (ACX) é outra importante fonte de H2O2 nos peroxissomos (BAKER et al., 2006; KAUR et al., 2009). Outra fonte desta ERO é o catabolismo de poliaminas (SANDALIO et al., 2013).

Além da catalase, as enzimas APX, MDAR, DHAR e GR também estão envolvidas na remoção do H2O2 gerado na fotorrespiração (MULLEN et al., 1999; KARYOTOU; DONALDSON, 2005; LISENBEE et al., 2005), por meio do ciclo ascorbato-glutationa (FOYER; NOCTOR, 2011). A distribuição das diferentes enzimas tem sido estudada em peroxissomos de folhas de ervilha, a APX foi localizada no lado citosólico da membrana peroxissomal (JIMÉNEZ et al. 1997), MDAR foi localizada na membrana e na matriz (LÓPEZ-HUERTAS et al., 1999; LETERRIER et al., 2005; LISENBEE et al., 2005), e GR apenas na matriz (JIMÉNEZ et al., 1997; ROMERO-PUERTAS et al., 2006).

A glutationa-S-transferase (GST), e as peroxiredoxinas (Prx) são outras enzimas antioxidantes presentes nos peroxissomos (Del RIO et al., 2006). A GST está envolvida na desintoxicação através da conjugação de glutationa a vários metabólitos eletrofílicos, direcionando-os para os vacúolos (EDWARDS et al., 2000). A atividade de GPX está relacionada com tolerância a estresse oxidativo. Deste modo, é provável que esta enzima ajude a anular a ameaça representada por alto nível de ERO e hidroperóxidos de ácidos graxos fitotóxicos presentes nos peroxissomos (CUMMINS et al., 1999; DIXON et al., 2009).

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