Rui Zhu , Tiesong Hu , Giulia Vico , Lu Zhang , Rangjian Qiu , Yingping Wang , Yong Li , Yong Liu
{"title":"气孔、叶肉和生化限制如何共同限制光合作用:一个双层优化模型","authors":"Rui Zhu , Tiesong Hu , Giulia Vico , Lu Zhang , Rangjian Qiu , Yingping Wang , Yong Li , Yong Liu","doi":"10.1016/j.agrformet.2025.110829","DOIUrl":null,"url":null,"abstract":"<div><div>Photosynthesis depends on stomatal, mesophyll, and biochemical limitations, which in turn respond to the environment in complex ways. However, it remains unclear how to quantitatively describe the interrelationships among these three limitations and their response to environmental changes.</div><div>We introduce a nested (bilevel) optimization modeling framework in which stomatal and non-stomatal limitations are optimized for separate optimization objectives. Stomatal optimization is prioritized, with stomatal conductance maximizing photosynthesis for set hydraulic costs. In turn, for set stomatal conductance, mesophyll conductance and biochemical capacity are optimized for maximum photosynthesis at a minimal cost.</div><div>Our model reproduced observed responses of stomatal conductance, mesophyll conductance, and biochemical capacity to key environmental factors (light, air <span><math><mrow><mi>C</mi><msub><mi>O</mi><mn>2</mn></msub></mrow></math></span> concentration, vapor pressure deficit, and soil moisture), including the different but coordinated reductions in stomatal conductance, mesophyll conductance, and biochemical capacity under water stress. Crucially, the model correctly estimated changes in intrinsic water use efficiency and provided testable predictions about how variations in plant hydraulic and photosynthetic traits drive the dynamic of photosynthetic limitations.</div><div>Our work provides a novel optimization framework for understanding and predicting how mesophyll and biochemical limitations are coordinated with stomatal regulation, which facilitates further theoretical and experimental studies.</div></div>","PeriodicalId":50839,"journal":{"name":"Agricultural and Forest Meteorology","volume":"375 ","pages":"Article 110829"},"PeriodicalIF":5.7000,"publicationDate":"2025-09-13","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":"0","resultStr":"{\"title\":\"How stomatal, mesophyll, and biochemical limitations co-limit photosynthesis: A bilevel optimization model\",\"authors\":\"Rui Zhu , Tiesong Hu , Giulia Vico , Lu Zhang , Rangjian Qiu , Yingping Wang , Yong Li , Yong Liu\",\"doi\":\"10.1016/j.agrformet.2025.110829\",\"DOIUrl\":null,\"url\":null,\"abstract\":\"<div><div>Photosynthesis depends on stomatal, mesophyll, and biochemical limitations, which in turn respond to the environment in complex ways. However, it remains unclear how to quantitatively describe the interrelationships among these three limitations and their response to environmental changes.</div><div>We introduce a nested (bilevel) optimization modeling framework in which stomatal and non-stomatal limitations are optimized for separate optimization objectives. Stomatal optimization is prioritized, with stomatal conductance maximizing photosynthesis for set hydraulic costs. In turn, for set stomatal conductance, mesophyll conductance and biochemical capacity are optimized for maximum photosynthesis at a minimal cost.</div><div>Our model reproduced observed responses of stomatal conductance, mesophyll conductance, and biochemical capacity to key environmental factors (light, air <span><math><mrow><mi>C</mi><msub><mi>O</mi><mn>2</mn></msub></mrow></math></span> concentration, vapor pressure deficit, and soil moisture), including the different but coordinated reductions in stomatal conductance, mesophyll conductance, and biochemical capacity under water stress. Crucially, the model correctly estimated changes in intrinsic water use efficiency and provided testable predictions about how variations in plant hydraulic and photosynthetic traits drive the dynamic of photosynthetic limitations.</div><div>Our work provides a novel optimization framework for understanding and predicting how mesophyll and biochemical limitations are coordinated with stomatal regulation, which facilitates further theoretical and experimental studies.</div></div>\",\"PeriodicalId\":50839,\"journal\":{\"name\":\"Agricultural and Forest Meteorology\",\"volume\":\"375 \",\"pages\":\"Article 110829\"},\"PeriodicalIF\":5.7000,\"publicationDate\":\"2025-09-13\",\"publicationTypes\":\"Journal Article\",\"fieldsOfStudy\":null,\"isOpenAccess\":false,\"openAccessPdf\":\"\",\"citationCount\":\"0\",\"resultStr\":null,\"platform\":\"Semanticscholar\",\"paperid\":null,\"PeriodicalName\":\"Agricultural and Forest Meteorology\",\"FirstCategoryId\":\"97\",\"ListUrlMain\":\"https://www.sciencedirect.com/science/article/pii/S0168192325004484\",\"RegionNum\":1,\"RegionCategory\":\"农林科学\",\"ArticlePicture\":[],\"TitleCN\":null,\"AbstractTextCN\":null,\"PMCID\":null,\"EPubDate\":\"\",\"PubModel\":\"\",\"JCR\":\"Q1\",\"JCRName\":\"AGRONOMY\",\"Score\":null,\"Total\":0}","platform":"Semanticscholar","paperid":null,"PeriodicalName":"Agricultural and Forest Meteorology","FirstCategoryId":"97","ListUrlMain":"https://www.sciencedirect.com/science/article/pii/S0168192325004484","RegionNum":1,"RegionCategory":"农林科学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":null,"EPubDate":"","PubModel":"","JCR":"Q1","JCRName":"AGRONOMY","Score":null,"Total":0}
How stomatal, mesophyll, and biochemical limitations co-limit photosynthesis: A bilevel optimization model
Photosynthesis depends on stomatal, mesophyll, and biochemical limitations, which in turn respond to the environment in complex ways. However, it remains unclear how to quantitatively describe the interrelationships among these three limitations and their response to environmental changes.
We introduce a nested (bilevel) optimization modeling framework in which stomatal and non-stomatal limitations are optimized for separate optimization objectives. Stomatal optimization is prioritized, with stomatal conductance maximizing photosynthesis for set hydraulic costs. In turn, for set stomatal conductance, mesophyll conductance and biochemical capacity are optimized for maximum photosynthesis at a minimal cost.
Our model reproduced observed responses of stomatal conductance, mesophyll conductance, and biochemical capacity to key environmental factors (light, air concentration, vapor pressure deficit, and soil moisture), including the different but coordinated reductions in stomatal conductance, mesophyll conductance, and biochemical capacity under water stress. Crucially, the model correctly estimated changes in intrinsic water use efficiency and provided testable predictions about how variations in plant hydraulic and photosynthetic traits drive the dynamic of photosynthetic limitations.
Our work provides a novel optimization framework for understanding and predicting how mesophyll and biochemical limitations are coordinated with stomatal regulation, which facilitates further theoretical and experimental studies.
期刊介绍:
Agricultural and Forest Meteorology is an international journal for the publication of original articles and reviews on the inter-relationship between meteorology, agriculture, forestry, and natural ecosystems. Emphasis is on basic and applied scientific research relevant to practical problems in the field of plant and soil sciences, ecology and biogeochemistry as affected by weather as well as climate variability and change. Theoretical models should be tested against experimental data. Articles must appeal to an international audience. Special issues devoted to single topics are also published.
Typical topics include canopy micrometeorology (e.g. canopy radiation transfer, turbulence near the ground, evapotranspiration, energy balance, fluxes of trace gases), micrometeorological instrumentation (e.g., sensors for trace gases, flux measurement instruments, radiation measurement techniques), aerobiology (e.g. the dispersion of pollen, spores, insects and pesticides), biometeorology (e.g. the effect of weather and climate on plant distribution, crop yield, water-use efficiency, and plant phenology), forest-fire/weather interactions, and feedbacks from vegetation to weather and the climate system.