马尾松幼苗光合产物的运输与分配特征

doi:10.11707/j.1001-7488.20190703

摘要/Abstract

摘要： [目的]解析光合产物在马尾松幼苗植株中的运输和分配规律，揭示马尾松生产力形成过程，为探讨不同环境干扰下马尾松光合产物分配过程的变化提供参考依据。[方法]采用¹³C同位素脉冲标记法，对1.5年生马尾松幼苗进行标记，标记结束后第0、2、5、17、24、72、120、168、216和360 h，按不同部位对马尾松幼苗进行全收获取样，测定¹³C含量，以监测近期合成的光合产物在马尾松幼苗中的运输和分配规律，同时测定光合产物全碳、非结构性碳水化合物（NSC）在各个器官的积累量。[结果]1）标记的光合产物在针叶中合成后，向各库器官的运输量随着时间延长由多逐渐减少，具体表现为标记结束后0~24 h内最多，24~216 h逐渐减少，216 h之后运输完成，且59%以上标记的光合产物在0~24 h内运输到各库器官。2）光合产物运输趋于稳定后，在各器官的分配大小依次为1年生叶 > 当年生叶 > 根 > 茎干 > 1年生枝 > 当年生枝，与生物量的分配大小一致，但与库活力大小不同，其库活力依次为当年生叶 > 当年生枝 > 1年生叶 > 根 > 1年生枝 > 茎干。3）各器官全碳和NSC积累量的分配与近期合成光合产物的分配大小一致，依次为1年生叶 > 当年生叶 > 根 > 茎干 > 1年生枝 > 当年生枝。[结论]马尾松幼苗光合产物运输速率大于0.1 m·h^-1，59%以上标记的光合产物在合成后的0~24 h内完成向各个库器官输出。新合成的光合产物在各器官中的积累量表现为功能器官（叶和根）居多，这一分配规律有利于马尾松幼苗阶段的生长。

关键词: 光合产物运输, 光合产物分配, 光合产物的积累, ¹³C脉冲标记, 马尾松

Abstract: [Objective] This study aimed to understand the pattern of transport and distribution of photosynthates in Pinus massoniana and the formation process of P. massoniana productivity.[Method] In this study, the 1.5-year-old P. massoniana seedlings were fed with pulse-chased ¹³C labeling for 4 h, and then the different organs were collected at 0, 2, 5, 17, 24, 72, 120, 168, 216 and 360 h to analyze the ¹³C levels, to investigate the transport and distribution of recently synthesized photosynthates in the seedlings. Meanwhile, the accumulation of photosynthates (total carbon and non-structural carbohydrate (NSC)) was determined in various organs.[Result] 1) After the labeled photosynthetic products were synthesized in conifers, the labeled photosynthates were found to be transported to the sink organs, however the amount of the transportation gradually decreased with time. It was manifested that most of the photosynthates were transported within 0-24 hours after the labeling, and the transportation was decreased gradually from 24 to 216 hours. In 216 hours, the labeled photosynthates finished outward transport. It was found that more than 59% of the labeled photosynthates were transported to the sink organs within 0-24 hours over the period of 216 hours. 2) After the transport of photosynthates tended to stable, the distribution of recently synthesized photosynthates in each organ was in the order of 1-year leaves > current-year leaves > roots > stem > 1-year branches > current-year branches, which was consistent with the distribution of the biomass, but different from the sink activity, the sink activity was ordered as current-year leaves > current-year branches > 1-year leaves > roots > 1-year branches > stem. 3) The distribution of total carbon and NSC accumulation in various organs was in accordance with the distribution of recently synthetic photosynthates, which was ordered as 1-year leaves > current-year leaves > roots > stem > 1-year branches > current-year branches.[Conclusion] The photosynthates transport rate in P. massoniana seedlings was greater than 0.1 m·h^-1, and more than 59% labeled photosynthates were exported to all sink organs within 0-24 hours after synthesis. The accumulation of new synthesized photosynthates in various organs was mostly in functional organs (leaves and roots). This distribution pattern is conducive to the growth of P. massoniana at seedling stage.

Key words: transport of photosynthates, distribution of photosynthates, accumulation of photosynthates, ¹³C pulse labelling, Pinus massoniana

中图分类号:

S718.43

邓秀秀, 肖文发, 曾立雄, 雷蕾, 施征. 马尾松幼苗光合产物的运输与分配特征[J]. 林业科学, 2019, 55(7): 27-34.

Deng Xiuxiu, Xiao Wenfa, Zeng Lixiong, Lei Lei, Shi Zheng. Transport and Distribution Characteristics of Photosynthates of Pinus massoniana Seedlings[J]. Scientia Silvae Sinicae, 2019, 55(7): 27-34.

参考文献

丁贵杰,周志春,王章荣,等. 2006.马尾松纸浆用材林培育与利用.北京:中国林业出版社, 1-34.
(Ding G J, Zhou Z C, Wang Z R, et al. 2006. Cultivation and utilization of pulpwood stand for Pinus massoniana. Beijing:China Forestry Publishing House, 1-34.[in Chinese])
杜明凤,丁贵杰,赵熙州. 2017.不同家系马尾松对持续干旱的响应及抗旱性.林业科学, 53(6):21-29.
(Du M F, Ding G J, Zhao X Z. 2017. Responses to continuous drought stress and drought resistance of different Masson Pine families. Scientia Silvae Sinicae, 53 (6):21-29.[in Chinese])
龚月桦,高俊凤,李锦辉. 1999.植物体内光合同化物韧皮部装载和卸出研究进展.西北植物学报, 19 (4):738-745.
(Gong Y H, Gao J F, Li J H. 1999. Progress in phloem loading and unloading of plant photoassimiate. Acta Botanica Boreali-Occidentalia Sinica, 19 (4):738-745.[in Chinese])
谷艳芳,丁圣彦,高志英,等. 2010.干旱胁迫下冬小麦光合产物分配格局及其与产量的关系.生态学报, 30 (5):1167-1173.
(Gu Y F, Ding S Y, Gao Z Y, et al. 2010. The pattern of photosynthate partitioning in drought-stressed winter wheat and its relationship with yield. Acta Ecologica Sinica, 30 (5):1167-1173.[in Chinese])
刘万德,苏建荣,李帅锋,等. 2017.云南普洱季风常绿阔叶林主要树种非结构性碳水化合物变异分析.林业科学, 53 (6):1-9.
(Liu W D, Su J R, Li S F, et al. 2017. Variation of non-structural carbohydrates for the dominant species in a monsoon broad-leaved evergreen forest in Pu'er, Yunnan Province. Scientia Silvae Sinicae, 53 (6):1-9.[in Chinese])
潘庆民,韩兴国,白永飞,等. 2002.植物非结构性贮藏碳水化合物的生理生态学研究进展.植物学通报, 19 (1):30-38.
(Pan Q M, Han X G, Bai Y F, et al. 2002. Advances in physiology and ecology studies on stored non-structure carbohydrates in plants. Chinese Bulletin of Botany, 19 (1):30-38.[in Chinese])
庞丽,张一,周志春,等. 2014.模拟氮沉降对低磷胁迫下马尾松生长和磷效率的影响.应用生态学报, 25 (5):1275-1282.
(Pang L, Zhang Y, Zhou Z C, et al. 2014. Effects of simulated nitrogen deposition on growth and phosphorus efficiency of Pinus massoniana under low phosphorus stress. Chinese Journal of Applied Ecology, 25(5):1275-1282.[in Chinese])
施征,白登忠,张维诚,等. 2017.青海云杉休眠前后非结构性碳水化合物含量随海拔变化.林业科学研究, 30 (6):908-915.
(Shi Z, Bai D Z, Zhang W C, et al. 2017.Variation of nonstructural carbohydrates (NSC) in Picea crassifolia at the alpine treeline of Qilian Mountains before and after dormancy. Forest Research, 30 (6):908-915.[in Chinese])
孙昭安,陈清,韩笑,等. 2018. ¹³C脉冲标记法定量冬小麦光合碳分配及其向地下的输入.环境科学, 39 (6):1-10.
(Sun S A, Chen Q, Han X, et al. 2018. Estimation of winter wheat photosynthesized carbon distribution and allocation belowground via ¹³C pulse-labeling. Environmental Science, 39 (6):1-10.[in Chinese])
宋纯鹏,王学路,周云,等. 2015.植物生理.北京:科学出版社, 179-234.
(Song C P, Wang X L, Zhou Y, et al. Plant physiology. 2015. Beijing:Science Press, 179-234.[in Chinese])
王云龙,许振柱,周广胜. 2004.水分胁迫对羊草光合产物分配及其气体交换特征的影响.植物生态学报, 28 (6):803-809.
(Wang Y L, Xu Z Z, Zhou G S. 2004. Changes in biomass allocation and gas exchange characteristics of Leymus chinensis in response to soil water stress. Acta Phytoecologica Sinica, 28 (6):803-809.[in Chinese])
韦莉莉,张小全,侯振宏,等. 2005.杉木苗木光合作用及其产物分配对水分胁迫的响应.植物生态学报, 29 (3):394-402.
(Wei L L, Zhang X Q, Hou Z H, et al. 2005. Effects of water stress on photosynthesis and carbon allocation in Cunninghamia lanceolata seedlings. Acta Phytoecologica Sinica, 29(3):394-402.[in Chinese])
杨世超,邓波,张蕴薇,等. 2012.中苜一号紫花苜蓿碳同化分配与转移速率的研究.草地学报, 20 (4):679-685.
(Yang S C, Deng B, Zhang Y W, et al. 2012. Absorption and transfer of photosynthetic carbon in Alfalfa (Zhongmu No.1). Acta Agrestia Sinica, 20 (4):679-685.[in Chinese])
张建国,盛炜彤,罗红艳,等. 2003. N、P、NP营养对杉木苗木生长和光合产物分配的影响.林业科学, 39 (2):21-27.
(Zhang J G, Sheng W T, Luo H Y, et al. 2003. Effects of N, P and NP nutrition on growth and allocation of photosynthetic products of Chinese fir seedlings. Scientia Silvae Sinicae, 39 (2):21-27.[in Chinese])
Adams H D, Germino M J, Breshears D D, et al. 2013. Nonstructural leaf carbohydrate dynamics of Pinus edulis during drought-induced tree mortality reveal role for carbon metabolism in mortality mechanism. New Phytologist, 197 (4):1142-1151.
Bahn M, Schmitt M, Siegwolf R, et al. 2009. Does photosynthesis affect grassland soil-respired CO₂ and its carbon isotope composition on a diurnal timescale?New Phytologist, 182 (2):451-460.
Blessing C H, Werner R A, Siegwolf R, et al. 2015. Allocation dynamics of recently fixed carbon in beech saplings in response to increased temperatures and drought. Tree Physiology, 35 (6):585-598.
Domisch T, Finér L, Lehto T. 2001. Effects of soil temperature on biomass and carbohydrate allocation in Scots pine (Pinus sylvestris) seedlings at the beginning of the growing season. Tree Physiology, 21 (7):465-472.
Hansen J, Beck E. 1990. The fate and path of assimilation products in the stem of 8-year-old Scots pine (Pinus sylvestris L.) trees. Trees, 4 (1):16-21.
Ho L C. 1988. Metabolism and compartmentation of imported sugars in sink organs in relation to sink strength. Annual Review of Plant Biology, 39 (1):355-378.
Koch K E. 1996. Carbohydrate-modulated gene expression in plants. Annual Review of Plant Physiology and Plant Molecular Biology, 47 (1):509-540.
Kozlowski T T. 1992. Carbohydrate sources and sinks in woody plants. Botanical Review, 58 (2):107-222.
Lacointe A. 2000. Carbon allocation among tree organs:a review of basic processes and representation in functional-structural tree models. Annals of Forest Science, 57 (5):521-533.
Lippu J. 1994. Patterns of dry matter partitioning and ¹⁴C-photosynthate allocation in 1.5-year-old Scots pine seedlings. Silva Fennica, 28 (3):145-153.
Myers J A, Kitajima K. 2007. Carbohydrate storage enhances seedling shade and stress tolerance in a Neotropical forest. Journal of Ecology, 95 (2):383-395.
Ngugi M R, Hunt M A, Doley D, et al. 2003. Dry matter production and allocation in Eucalyptus cloeziana and Eucalyptus argophloia seedlings in response to soil water deficits. New Forests, 26 (2):187-200.
Pumpanen J S, Heinonsalo J, Rasilo T, et al. 2009. Carbon balance and allocation of assimilated CO₂ in Scots pine, Norway spruce, and Silver birch seedlings determined with gas exchange measurements and ¹⁴C pulse labelling. Trees, 23 (3):611-621.
Schneider A, Schmitz K. 1989. Seasonal course of translocation and distribution of ¹⁴C-labelled photoassimilate in young trees of Larix decidua Mill. Trees, 3 (4):185-191.
Sun F, Kuang Y, Wen D, et al. 2010. Long-term tree growth rate, water use efficiency, and tree ring nitrogen isotope composition of Pinus massoniana L. in response to global climate change and local nitrogen deposition in Southern China. Journal of Soils and Sediments, 10 (8):1453-1465.
Thompson R G, Fensom D S, Anderson R R, et al. 1979. Translocation of ¹¹C from leaves of Helianthus, Heracleum, Nymphoides, Ipomoea, Tropaeolum, Zea, Fraxinus, Ulmus, Picea, and Pinus: comparative shapes and some fine structure profiles. Canadian Journal of Botany, 57 (8):845-863.
Wang G, Liu F. 2014. Carbon allocation of Chinese pine seedlings along a nitrogen addition gradient. Forest Ecology and Management, 334:114-121.
Wiley E, Huepenbecker S, Casper B B, et al. 2013. The effects of defoliation on carbon allocation:Can carbon limitation reduce growth in favour of storage? Tree Physiology, 33 (11):1216-1228.
Zang U, Goisser M, Grams T E, et al. 2014. Fate of recently fixed carbon in European beech (Fagus sylvatica) saplings during drought and subsequent recovery. Tree Physiology, 34 (1):29-38.
Zhang Y, Zhou Z, Yang Q. 2013. Nitrogen (N) deposition impacts seedling growth of Pinus massoniana via N:P ratio effects and the modulation of adaptive responses to low P (Phosphorus). PloS One, 8 (10):1-9.
Zimmermann M H. 1958. Translocation of organic substances in the phloem of trees//The physiology of forest trees. New York:Ronald Press, 381-400.

[1]	叶钰倩, 赵家豪, 刘畅, 关庆伟. 间伐强度对马尾松人工林根际与非根际土壤中性糖特征的影响[J]. 林业科学, 2019, 55(8): 28-35.
[2]	金国庆, 张振, 余启新, 冯随起, 丰忠平, 赵世荣, 周志春. 马尾松2个世代种子园6年生家系生长的遗传变异与增益比较[J]. 林业科学, 2019, 55(7): 57-67.
[3]	李亚藏, 冯仲科. 气候敏感的马尾松生物量相容性方程系统研建[J]. 林业科学, 2019, 55(5): 65-73.
[4]	王好运, 吴峰, 朱小坤, 谢维斌. 叶型对马尾松幼苗生长及叶绿素荧光特征的影响[J]. 林业科学, 2019, 55(3): 183-192.
[5]	陈金磊, 方晰, 辜翔, 李雷达, 刘兆丹, 王留芳, 张仕吉. 中亚热带2种森林群落组成、结构及区系特征[J]. 林业科学, 2019, 55(2): 159-172.
[6]	陈坦, 张振, 楚秀丽, 金国庆, 周志春, 丰忠平. 马尾松二代无性系种子园的花期同步性[J]. 林业科学, 2019, 55(1): 146-156.
[7]	罗晓蔓, 丁贵杰, 王艺. 菌根化马尾松苗木根际土壤浸提物有效成份及其对种子萌发的影响[J]. 林业科学, 2018, 54(8): 32-38.
[8]	姚虹宇, 刘亚敏, 张盛楠, 刘玉民, 周文颖, 王针针. 外源柠檬酸对铝胁迫下马尾松生理特性的影响[J]. 林业科学, 2018, 54(7): 155-164.
[9]	魏永成, 刘青华, 周志春, 徐六一, 陈雪莲, 郝焰平. 不同抗性马尾松接种松材线虫后针叶内化学信号物质的变化[J]. 林业科学, 2018, 54(2): 119-125.
[10]	辜夕容, 倪亚兰, 江亚男, 贾豪. 接种双色蜡蘑对酸性铝胁迫下马尾松幼苗生长、养分和铝吸收与分布的影响[J]. 林业科学, 2018, 54(2): 170-178.
[11]	尹焕焕, 刘青华, 周志春, 余启新, 丰忠平. 马尾松产脂性状与生长性状的无性系变异及相关性[J]. 林业科学, 2018, 54(12): 82-91.
[12]	高尚坤, 肖文发, 曾立雄, 雷蕾, 黄志霖, 王松. 马尾松人工林干扰对土壤微生物群落结构的短期影响[J]. 林业科学, 2018, 54(12): 92-101.
[13]	张振, 金国庆, 余启新, 刘青华, 丰忠平, 董虹妤, 周志春. 马尾松三代测定林抽梢性状的遗传效应及与针叶氮磷钾养分的遗传相关[J]. 林业科学, 2017, 53(8): 26-34.
[14]	杜明凤, 丁贵杰, 赵熙州. 不同家系马尾松对持续干旱的响应及抗旱性[J]. 林业科学, 2017, 53(6): 21-29.
[15]	贺同鑫, 孙建飞, 李艳鹏, 俞有志, 胡宝清, 王清奎. 环割对杉木和马尾松人工林土壤微生物群落结构的影响[J]. 林业科学, 2017, 53(6): 77-84.