van Dongen Joost T, Roeb Gerhard W, Dautzenberg Marco, Froehlich Anja, Vigeolas Helene, Minchin Peter E H, Geigenberger Peter
Max Planck Institute of Molecular Plant Physiology, Am Mühlenberg 1, 14476 Golm, Germany.
Plant Physiol. 2004 Jul;135(3):1809-21. doi: 10.1104/pp.104.040980. Epub 2004 Jul 9.
We studied the influence of the internal oxygen concentration in seeds of wheat (Triticum aestivum) on storage metabolism and its relation to phloem import of nutrients. Wheat seeds that were developing at ambient oxygen (21%) were found to be hypoxic (2.1%). Altering the oxygen supply by decreasing or increasing the external oxygen concentration induced parallel changes in the internal oxygen tension. However, the decrease in internal concentration was proportionally less than the reduction in external oxygen. This indicates that decreasing the oxygen supply induces short-term adaptive responses to reduce oxygen consumption of the seeds. When external oxygen was decreased to 8%, internal oxygen decreased to approximately 0.5% leading to a decrease in energy production via respiration. Conversely, increasing the external oxygen concentration above ambient levels increased the oxygen content as well as the energy status of the seeds, indicating that under normal conditions the oxygen supply is strongly limiting for energy metabolism in developing wheat seeds. The intermediate metabolites of seed storage metabolism were not substantially affected when oxygen was either increased or decreased. However, at subambient external oxygen concentrations (8%) the metabolic flux of carbon into starch and protein, measured by injecting (14)C-Suc into the seeds, was reduced by 17% and 32%, respectively, whereas no significant effect was observed at superambient (40%) oxygen. The observed decrease in biosynthetic fluxes to storage compounds is suggested to be part of an adaptive response to reduce energy consumption preventing excessive oxygen consumption when oxygen supply is limited. Phloem transport toward ears exposed to low (8%) oxygen was significantly reduced within 1 h, whereas exposing ears to elevated oxygen (40%) had no significant effect. This contrasts with the situation where the distribution of assimilates has been modified by removing the lower source leaves from the plant, resulting in less assimilates transported to the ear in favor of transport to the lower parts of the plant. Under these conditions, with two strongly competing sinks, elevated oxygen (40%) did lead to a strong increase in phloem transport to the ear. The results show that sink metabolism is affected by the prevailing low oxygen concentrations in developing wheat seeds, determining the import rate of assimilates via the phloem.
我们研究了小麦(Triticum aestivum)种子内部氧气浓度对储存代谢的影响及其与韧皮部养分输入的关系。发现在环境氧气(21%)条件下发育的小麦种子处于低氧状态(2.1%)。通过降低或增加外部氧气浓度来改变氧气供应,会引起内部氧气张力的平行变化。然而,内部浓度的降低与外部氧气的减少相比,比例上要小。这表明降低氧气供应会引发短期适应性反应,以减少种子的氧气消耗。当外部氧气降至8%时,内部氧气降至约0.5%,导致通过呼吸作用产生的能量减少。相反,将外部氧气浓度提高到环境水平以上,会增加种子的氧气含量以及能量状态,这表明在正常条件下,氧气供应对发育中的小麦种子的能量代谢有很强的限制作用。当氧气增加或减少时,种子储存代谢的中间代谢产物没有受到实质性影响。然而,在低于环境的外部氧气浓度(8%)下,通过向种子中注射(14)C-蔗糖来测量的碳进入淀粉和蛋白质的代谢通量分别降低了17%和32%,而在高于环境的(40%)氧气浓度下未观察到显著影响。观察到的向储存化合物的生物合成通量的降低被认为是一种适应性反应的一部分,以减少能量消耗,防止在氧气供应有限时过度消耗氧气。暴露于低(8%)氧气环境下的麦穗,韧皮部运输在1小时内显著减少,而将麦穗暴露于高氧气(40%)环境下则没有显著影响。这与通过去除植物下部源叶来改变同化物分配的情况形成对比,在这种情况下,运往麦穗的同化物减少,有利于运往植物下部。在这些条件下,有两个强烈竞争的库,高氧气(40%)确实导致韧皮部向麦穗的运输大幅增加。结果表明,库代谢受到发育中小麦种子中普遍存在的低氧浓度的影响,决定了通过韧皮部的同化物输入速率。