Acta Physiologiae Plantarum

, 38:224 | Cite as

Exogenous spermidine delays chlorophyll metabolism in cucumber leaves (Cucumis sativus L.) under high temperature stress

  • Heng Zhou
  • Shirong Guo
  • Yahong An
  • Xi Shan
  • Ying Wang
  • Sheng Shu
  • Jin Sun
Original Article


This study evaluated the effects of exogenous spermidine (Spd) on chlorophyll (Chl) biosynthesis and catabolism in the leaves of cucumber (Cucumis sativus L.) seedlings under high temperature stress. Substrate culture experiments were performed using the high temperature-sensitive variety ‘Jinchun No. 2’ in an artificial climate chamber at 42/32 °C with foliar applications of 1.0 mmol L−1 Spd. The results suggested that high temperature stress markedly reduced the leaf Chl concentrations and inhibited plant growth; the harmful effect of high temperature on the cucumber seedlings was mitigated by exogenous Spd, which increased the leaf Chl concentration and promoted plant growth. Under high temperature stress, the conversion of porphobilinogen (PBG) into uroporphyrinogen III (UroIII) in the Chl biosynthetic pathway and the catabolic process of Chl were accelerated. Following the application of exogenous Spd, the conversion of PBG into UroIII was suppressed, and the accumulation of certain intermediates, e.g., protoporphyrin IX (ProtoIX) and Mg-protoporphyrin IX (Mg-ProtoIX), was decreased in the Chl biosynthetic pathway. Additionally, exogenous Spd reduced chlorophyllase (Chlase) and Mg-dechelatase (MDCase) activity and transcript levels and markedly downregulated pheophorbide A oxygenase (PaO), red Chl catabolite reductase (RCCR), Chl b reductase 1 (CBR1) and stay-green reductase 1 (SGR1) transcript levels. These results indicate that although high temperature stress accelerated Chl biosynthesis, it concurrently facilitated Chl catabolism in cucumber leaves. Exogenous Spd delayed the conversion of PBG into UroIII in the Chl biosynthetic pathway, effectively preventing oxidative bleaching of Chl in cucumber leaves. Meanwhile, Spd clearly decreased PaO pathway-related enzyme activity and transcript levels, thereby slowing Chl catabolism and increasing Chl concentrations.


High temperature stress Spermidine Chlorophyll Biosynthesis Degradation 



δ-Aminolevulinic acid


Chl b reductase


Chl catabolic enzymes





Chlide a

Chlorophyllide a




Hydroxymethylbilane synthase




Mg-protoporphyrin IX


Nonfluorescent Chl catabolites


Photosystem II


Chl–protein complex


Primary fluorescent Chl catabolites




Pheophorbide A oxygenase




Porphobilinogen deaminase


Proto Chl

Pheide a

Pheophorbide a

Pheide b

Pheophorbide b


Protoporphyrin IX


Red Chl catabolite reductase


Red Chl catabolites


Reactive oxygen species


Stay-green reductase 1




Uroporphyrinogen III


Uroporphyrinogen III synthase



This work was supported by the National Natural Science Foundation of China (Nos. 31471869, 31401919, and 31272209), the Central Research Institutes of Basic Research Fund (6J0745), the Priority Academic Program Development of Jiangsu Higher Education Institutions (PDPA), and the China Agriculture Research System (CARS-25-C-03) and was sponsored by the Research Fund for the Doctoral Program of Higher Education (20130097120015).

Compliance with ethical standards

Conflict of interest

The authors have no conflicts of interest to declare.


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Copyright information

© Franciszek Górski Institute of Plant Physiology, Polish Academy of Sciences, Kraków 2016

Authors and Affiliations

  • Heng Zhou
    • 1
    • 2
  • Shirong Guo
    • 1
    • 2
  • Yahong An
    • 1
    • 2
  • Xi Shan
    • 1
    • 2
  • Ying Wang
    • 1
    • 2
  • Sheng Shu
    • 1
    • 2
  • Jin Sun
    • 1
    • 2
  1. 1.College of HorticultureNanjing Agriculture UniversityNanjingPeople’s Republic of China
  2. 2.Jiangsu Province Engineering Laboratory for Modern Facility Agriculture Technology and EquipmentNanjingPeople’s Republic of China

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