Ilík Petr, Schansker Gert, Kotabová Eva, Váczi Peter, Strasser Reto J, Barták Milos
Laboratory of Biophysics, Faculty of Science, Palacký University, tr. Svobody 26, CZ-77146 Olomouc, Czech Republic.
Biochim Biophys Acta. 2006 Jan;1757(1):12-20. doi: 10.1016/j.bbabio.2005.11.008. Epub 2005 Dec 13.
An unusual dip (compared to higher plant behaviour under comparable light conditions) in chlorophyll fluorescence induction (FI) at about 0.2-2 s was observed for thalli of several lichen species having Trebouxia species (the most common symbiotic green algae) as their native photobionts and for Trebouxia species cultured separately in nutrient solution. This dip appears after the usual O(J)IP transient at a wide range of excitation light intensities (100-1800 micromol photons m(-2) s(-1)). Simultaneous measurements of FI and 820-nm transmission kinetics (I(820)) with lichen thalli showed that the decreasing part of the fluorescence dip (0.2-0.4 s) is accompanied by a decrease of I(820), i.e., by a reoxidation of electron carriers at photosystem I (PSI), while the subsequent increasing part (0.4-2 s) of the dip is not paralleled by the change in I(820). These results were compared with that measured with pea leaves-representatives of higher plants. In pea, PSI started to reoxidize after 2-s excitation. The simultaneous measurements performed with thalli treated with methylviologen (MV), an efficient electron acceptor from PSI, revealed that the narrow P peak in FI of Trebouxia-possessing lichens (i.e., the I-P-dip phase) gradually disappeared with prolonged MV treatment. Thus, the P peak behaves in a similar way as in higher plants where it reflects a traffic jam of electrons induced by a transient block at the acceptor side of PSI. The increasing part of the dip in FI remained unaffected by the addition of MV. We have found that the fluorescence dip is insensitive to antimycin A, rotenone (inhibitors of cyclic electron flow around PSI), and propyl gallate (an inhibitor of plastid terminal oxidase). The 2-h treatment with 5 microM nigericin, an ionophore effectively dissipating the pH-gradient across the thylakoid membrane, did not lead to significant changes either in FI nor I(820) kinetics. On the basis of the presented results, we suggest that the decreasing part of the fluorescence dip in FI of Trebouxia-lichens reflects the activation of ferredoxin-NADP(+)-oxidoreductase or Mehler-peroxidase reaction leading to the fast reoxidation of electron carriers in thylakoid membranes. The increasing part of the dip probably reflects a transient reduction of plastoquinone (PQ) pool that is not associated with cyclic electron flow around PSI. Possible causes of this MV-insensitive PQ reduction are discussed.
在几种以共球藻属(最常见的共生绿藻)为天然光共生体的地衣物种的叶状体以及在营养液中单独培养的共球藻属物种中,观察到叶绿素荧光诱导(FI)在约0.2 - 2秒处出现异常下降(与在可比光照条件下高等植物的行为相比)。这种下降在通常的O(J)IP瞬变之后出现,且在很宽的激发光强度范围内(100 - 1800微摩尔光子·米⁻²·秒⁻¹)都存在。用地衣叶状体同时测量FI和820纳米透射动力学(I(820))表明,荧光下降的下降部分(0.2 - 0.4秒)伴随着I(820)的下降,即光系统I(PSI)处电子载体的再氧化,而下降随后的上升部分(0.4 - 2秒)与I(820)的变化并不平行。这些结果与用豌豆叶(高等植物的代表)测量的结果进行了比较。在豌豆中,PSI在2秒激发后开始再氧化。用甲基紫精(MV,一种来自PSI的有效电子受体)处理叶状体后进行的同时测量表明,含有共球藻属的地衣FI中的窄P峰(即I - P下降阶段)随着MV处理时间延长逐渐消失。因此,P峰的行为与高等植物中相似,在高等植物中它反映了PSI受体侧瞬时阻断诱导的电子拥堵。FI下降的上升部分不受MV添加的影响。我们发现荧光下降对抗霉素A、鱼藤酮(PSI周围循环电子流的抑制剂)和没食子酸丙酯(质体末端氧化酶的抑制剂)不敏感。用5微摩尔尼日利亚菌素(一种有效消散类囊体膜跨膜pH梯度的离子载体)处理2小时,无论是FI还是I(820)动力学都没有导致显著变化。基于所呈现的结果,我们认为共球藻属地衣FI中荧光下降的下降部分反映了铁氧还蛋白 - NADP⁺氧化还原酶或梅勒过氧化物酶反应的激活,导致类囊体膜中电子载体的快速再氧化。下降的上升部分可能反映了质体醌(PQ)库的瞬时还原,这与PSI周围的循环电子流无关。讨论了这种对MV不敏感的PQ还原的可能原因。