carbon cycle in plant canopyatmenv.envi.osakafu-u.ac.jp/.../251/ecolmet.2018_ppt2.pdfecosystem...
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生態気象学特論
Carbon Cycle in Plant Canopy
Ecosystem process and modeling
生態気象・植山
Ecosystem Processes
Examples : 光合成、呼吸、分解蒸発、蒸散、昇華、流出成長、分配、脱窒
Material Cycle:Water, Carbon, Water, and Energy
Compartment
Leaf
Stem
RootLitter
Soil
(Pool)
FluxPhotosynthesis
Respiration
decomposition
decompositiondecomposition litterfall
mortality
Leaf sprout
Retranslocation
Respiration
Carbon
Flux
PrecipitationSoilevaporation
water
Outflow
Transpiration
Intercepted evaporation
Precipitation
Snow
sublimation
Snow melt
Flux
deposition
nitrification
denitrification Litterfall
mortality
Leaf sprout
retranslocation
immobilization
nitrogen
(N-fixation)
Compartment Model
3211 FFF
t
C
C1 : Compartment1(e.g., leaf, stem, or root) (kg m-3)
t : Time (day)
Fi : Flux among compartments (e.g., photosynthesis, respiration, or litterfall) (kg m-2 day-1)
Explaining ecosystem processesusing Compartment and Flux
Carbon Flux
NPP = GPP – RaNEP = NPP – RhNEE = -NEP NBP = NEP + FCO + FCH4 + FVOC
+ FDIC + FDOC + FPOC - FFire
(Chapin et al., 2006)
NPP = GPP – Ra
(Chapin et al., 2006)
NPP : Net Primary Productivity純一次生産量
GPP : Gross Primary Productivity総一次生産量
Ra : Autotrophic respiration独立栄養呼吸
Carbon Flux
NPP (純一次生産量)
NPP = GPP – Ra
Slope = 0.53
CUE = NPP / GPPCarbon Use Efficiency (炭素利用効率)
(De Lucia et al., 2007)
森林について
NEP = NPP – Rh
(Chapin et al., 2006)
NPP : Net Primary Productivity純一次生産量
NEP : Net Ecosystem Productivity純生態系生産量
Rh : Heterotrophic respiration従属栄養呼吸
Carbon Flux
NEE = -NEP
(Chapin et al., 2006)
NEE : Net Ecosystem Exchange正味生態系交換量
NEP : Net Ecosystem Productivity純生態系生産量
Carbon Flux
NBP = NEP + FCO + FCH4 + FVOC
+ FDIC + FDOC + FPOC - FFire
(Chapin et al., 2006)
NBP : Net Biosphere Productivity正味生物圏交換量
F : Flux (フラックス)CO : 一酸化炭素 CH4 : メタン VOC : 揮発性有機物DIC : 溶存無機炭素 DOC : 溶存有機炭素 POC :粒子性有機炭素Fire : 火災 (その他の撹乱)
Carbon Flux
http://www.ipcc.ch/ipccreports/sres/land_use/index.php?idp=24
Carbon Flux
Photosynthesis
Stroma
Thylakoid(膜内にクロロフィル)
Grana
Photosynthetically active radiation:400~700 nm
Chloroplast photochemical reaction
Calvin cycle
(leaf-scale)
葉温
(Chapin et al., 2011)
チラコイド
ストロマ
(leaf-scale)
NADP (ニコチンアミドアデニンジヌクレオチドリン酸)
酵素ルビスコ触媒の酵素反応
クロロフィルの励起による
NA
DP
Hの生成
ホスホグリセリン酸
カルボキシル化
酸素化(光呼吸)
Photosynthesis
RuBP(リブロース・1,5-ビスリン酸)
(Chapin et al., 2011)
Photosynthesis
(Chapin et al., 2011)
(光が十分な場合)
Photosynthesis
A = min (Aj, Ac)
(彦坂, 2016)
Photosynthesis
Ci = 270 umol m-2 s-1
(Bernacchi et al., 2013)
A = min (Aj, Ac)Photosynthesis
Gas diffusion A = gt * (Ca - Cm)
Ca
Cm
(Chapin et al., 2011)
A = 1/rt * (Ca - Cm)
Cm
Ci
Cl
Ca
rm
rs
rb
rt = rb + rs + rm
A = 1/rm * (Ci - Cm)
A = 1/rs * (Cl - Ci)
A = 1/rb * (Ca - Cl)
Continuum
Gas diffusion
A = gt * (Ca - Cm)
Cm
Ci
Cl
Ca
rm
rs
rb
g = 1 / RA = gm * (Ci - Cm)
A = gs * (Cl - Ci)
A = gb * (Ca - Cl)
コンダクタンスは抵抗の逆数
Gas diffusion
Boundary-Layer Conductance (gb):葉の形状、風速 etc
Stomatal Conductance (gs):気孔の数、開度
Mesophyll conductance (gm):葉の厚さ、水分条件 etc
Gas diffusion
気孔コンダクタンス
蒸散による水損失を低減CO2取り込みによる生産の最大化
Hydropassive
Hydroactive
Gas diffusion
WUE (water use efficiency)水利用効率 = 光合成 / 蒸散
大気飽差 (kPa)
(米国;老齢林)
(Tang et al., 2006)
1gのCO2獲得に227gの水の損失
Gas diffusion
Modeling Stomatal Conductance
(Empirical)
Jarvis型モデル
(Running and Warning, 1998)
Jarvis型モデル
gsw = gsw_max * f(T) * f (VPD) * f (PAR)
* f (CO2) * f (SWC)
(Bond-Lamberty et al. 2007)
←潅水の経過日数によって気孔が決まるような関数
最大気孔開度のときの気孔コンダクタンス
気孔開度 (0~1)
Modeling Stomatal Conductance
gsw = A hs / Ca * m + b
光合成速度相対湿度
大気CO2濃度
経験定数 (パラメータ)
Ball型モデル
(Monson and Baldocchi, 2014)
Modeling Stomatal Conductance
(Empirical)
CO2濃度上昇で生態系プロセスはどう変化するか?
光合成速度
気孔コンダクタンス
蒸発散
土壌水分 窒素制限?
↑
↓
↓
↑
(von Caemmerer et al. 2009)
Photosynthesis - Condutance
https://nicholas.duke.edu/duke-forest-face
FACE (Free-Air Carbon Dioxide Enrichment)
開放系大気CO2濃度増加実験
(Long et al., 2004)
FACE (Free-Air Carbon Dioxide Enrichment)
開放系大気CO2濃度増加実験
0 60%CO2増加による光合成の変化率(%)
Photosynthesis vs Nitrogen content
(Chapin et al., 2011)
Canopy photosynthesis
PPFD (μmol m-2 s-1)
群落光合成 = 個葉光合成 x 葉面積 ??
(de Pury and Farquhar, 1997)
Radiation transfer - photosynthesisin plant canopy
Direct Diffuse
Radiation transfer - photosynthesisin plant canopy
Direct beam
Diffuse beam(Rayleigh scattering)
Direct beam
Diffuse beam(Mie scattering)
Radiation transfer - photosynthesis
sunfleck
in plant canopy
Radiation transfer - photosynthesisin plant canopy
PPFDsun = PPFDdirect + PPFDdiffuse + PPFDscatter
PPFDshade = PPFDdiffuse + PPFDscatter
Radiation at sunlit leaf
Radiation at shaded leaf
Radiation transfer - photosynthesisin plant canopy
6
葉面積指数 (m2 m-2)
実際に近い条件
(de Pury and Farquhar, 1997)
Radiation transfer – photosynthesis in plant canopy
Carbon flux
NPP = GPP – Ra
(Chapin et al., 2006)
NPP : Net Primary Productivity純一次生産量
GPP : Gross Primary Productivity総一次生産量
Ra : Autotrophic respiration独立栄養呼吸
Autotrophic respiration
Ra = Rg + Rm
Ra : Autotrophic respiration独立栄養呼吸
Rg : Growth respiration成長呼吸
Rm : Maintenance respiration維持呼吸
1 gの乾物を生成する際のコスト
(Chapin et al., 2011)
Autotrophic respiration
Maintenancerespiration
Growth respiration
Seasonality in autotrophic respiration at a temperate forest
(Miyama et al., 2006)
Autotrophic respiration
NPP (純一次生産量)
NPP = GPP – Ra
Slope = 0.53
CUE = NPP / GPPCarbon Use Efficiency (炭素利用効率)
(De Lucia et al., 2007)
NPP (純一次生産量)
NPP = GPP – Ra CUE = NPP / GPP
(De Lucia et al., 2007)
NPP (純一次生産量)
NPP = GPP – Ra (Chapin et al., 2011)
純一次生産量
https://nelson.wisc.edu/sage/data-and-models/atlas/maps.php?catnum=3&type=Ecosystems
各要素へのNPPの分配率の代表値
(Chapin et al., 2011)
Allocation(分配)
http://matome.naver.jp/odai/2143120929338037401
Allocation(分配)リービッヒの最小律
光に制限
地上部へ多く配分
水・栄養分に制限
地下部へ多く配分
Allocation(分配)
分配
(Littonet al., 2007GCB)
木部
地下部
葉
Carbon Flux
NEP = NPP – Rh
(Chapin et al., 2006)
NPP : Net Primary Productivity純一次生産量
NEP : Net Ecosystem Productivity純生態系生産量
Rh : Heterotrophic respiration従属栄養呼吸
NEP = NPP – Rh
(Chapin et al., 2011)
Decomposition
Litter
Litter
Litter
Litter
Soil
Soil
Soil
Soil
粗大有機物
Decomposition
CWD
*CWD : coarse woody debris
リター中の含水率(%)
従属栄養呼吸
(mg
CO
2g-1
liite
rd
-1)
(O’Connell et al., 1990)Decomposition
Carbon flux
(Ueyama al., 2009)Black spruce forest in Alaska
Methane budget in terrestrial ecosystems
Ecosystem process and modeling
GWP = 28 at 100 years
Greenhouse gas & radiative forcing
(IPCC AR5, 2013)
2nd Greenhouse Gas
http://www.esrl.noaa.gov/gmd/ccgg/iadv/C
H4
co
ncen
trati
on
(p
pb
)
Mauna Loa, Hawaii, USA
1700
1750
1800
1850
1900
1950
1650
1600 1984 1988 1992 1996 2000 2004 2008 2012
Year 2016
Methane
Pre-industrial level
722 ppb
678 Tg CH4 yr-1Methane emission
(Kirshke et al. 2013; Nature Geo.)
wetland
Agriculture
Fossil fuelBiomassburning
Wild animalsTermites
Others
Methane emission 678 Tg CH4 yr-1
(Kirshke et al. 2013; Nature Geo.)
wetland
Agriculture
Fossil fuelBiomassburning
Wild animalsTermites
Others
(Kirshke et al. 2013; Nature Geo.)
Methanogenic Archaea
632 Tg CH4 yr-1
OH-radical(83%)
Soil
Methane sinkStratospheric OH
Tropospheric
Methanotroph(bacteria)
(McEwing et al., 2015l Plant Soil)
Methane process in wetland
(Riley et al., 2011; Biogeosciences)
(Rinne et al., 2007; Tellus B)
Seasonality Boreal fen at Finland
Methane emission vs water table
(Oleefeldt et al., 2013; GCB)
High emissions
WetDry
Norther wetland across permafrost zones
Methane emission at wetlands
(Riley et al., 2011; Biogeosciences)
Modeling Global methane emission
(Melton et al., 2013; Biogeosciences)
Wetland area
(Melton et al., 2013; Biogeosciences)