Glacial CO2 Research Articles

Role of Southern Ocean stratification in glacial atmospheric CO2 reduction evaluated by a three-dimensional ocean general circulation model

Atmospheric carbon dioxide (CO2) concentration during glacial periods is known to be considerably lower than during interglacial periods. However, previous studies using an ocean general circulation model (OGCM) fail to reproduce this. Paleoclimate proxy data of the Last Glacial Maximum indicate high salinity (>37.0 psu) and long water mass residence time (>3,000 years) in the Southern Ocean, suggesting that salinity stratification was enhanced and more carbon was stored there. Reproducibility of salinity and water mass age is considered insufficient in previous OGCMs, which might affect the reproducibility of atmospheric CO2 concentration. This study investigated the role of increased stratification of the Southern Ocean in the glacial CO2 variation using an OGCM. We found that deep water formation in East Antarctica is required to explain high salinity in the South Atlantic. Saltier deep Southern Ocean resulted in increased atmospheric CO2 concentration against previous estimates. This is partly due to increased volume transport of Antarctic Bottom Water and associated decrease in the water mass age of the deep Pacific Ocean. On the other hand, weakening of vertical mixing contributed to increase of the vertical gradient of dissolved inorganic carbon and decrease of atmospheric CO2 concentration. However, we show that it is unable to explain all of the glacial CO2 variations by the contribution of the Southern Ocean. Our findings indicate that detailed understanding of the impact of enhanced stratification in the Southern Ocean on the Pacific Ocean might be crucial to understanding the mechanisms behind the glacial-interglacial ocean carbon cycle variations.

Two contributors to the glacial CO2 decline

It is generally accepted that the glacial drawdown of atmospheric CO2 content is the sole result of uptake by the ocean. Here we make a case that the reduction of planetary CO2 outgassing made a significant contribution. We propose that the ocean contribution to CO2 reduction closely followed Northern Hemisphere summer insolation and was superimposed on a ramp-like decline resulting from a reduction in the input of planetary CO2. We base this scenario on new records of δ13C and B to Ca ratio in cores from the upper and lower portions of the deep Atlantic. They demonstrate that the waxing and waning of the stratification of Atlantic deep water follows summer insolation. Our thoughts were driven by the observation that over the last 30 kyr the extent of mountain glaciation in both hemispheres appears to have tracked the atmosphere's CO2 content, suggesting that the connection between orbital cycles and land ice cover is via the ocean. Instead of a direct connection between ice extent and summer insolation, the tie is a modulation of the heat and fresh water budgets of the northern Atlantic. Changes in the boundary conditions lead to reorganizations of ocean circulation and, as a consequence, changes in CO2 storage in the ocean.

Elucidation of colour development and microstructural characteristics ofAllium sativumfumigated with acetic acid

Summary It has been well known that the greening of Allium sativum cloves could be formed after immersed in acetic acid solution. Nonetheless, no investigation was reported on colour development of A. sativum in response to acetic acid vapour until now. In this study, the brief exposure of A. sativum to acetic acid vapour (200–400 ppm) was combined with controlled atmosphere (5%, 20% and 80% CO2) packaging storage to break cell membrane and green garlic. The garlic bulbs were fumigated with acetic acid before controlled CO2 atmosphere packaging for 25 days at 4 °C. Fumigation with 200 ppm acetic acid followed by high CO2 atmosphere packaging (80% CO2) facilitated the garlic greening. It was also verified that γ-glutamyl transpeptidase was involved in garlic greening in present study, and the compromise of glacial acetic acid vapour fumigation and CO2 gas atmosphere in package stored at low temperature could result in garlic greening as well.

Glacial–interglacial changes in bottom-water oxygen content on the Portuguese margin

Some of the glacial CO2 drawdown has been attributed to CO2 storage in the deep Pacific and Southern oceans. Reconstruction of apparent oxygen utilization suggests that respired CO2 storage was also enhanced in the deep northeast Atlantic.

Responses of high-elevation herbaceous plant assemblages to low glacial CO₂ concentrations revealed by fossil marmot (Marmota) teeth.

Atmospheric CO2 cycles of the Quaternary likely imposed major constraints on the physiology and growth of C3 plants worldwide. However, the measured record of this remains both geographically and taxonomically sparse. We present the first reconstruction of physiological responses in a late Quaternary high-elevation herbaceous plant community from the Southern Rocky Mountains, USA. We used a novel proxy-fossilized tooth enamel of yellow-bellied marmots (Marmota flaviventris)-which we developed using detailed isotopic analysis of modern individuals. Calculated C isotopic discrimination (Δ) of alpine plants was nearly 2 ‰ lower prior to the Last Glacial Maximum than at present, a response almost identical to that of nonherbaceous taxa from lower elevations. However, initial shifts in Δ aligned most closely with the onset of the late Pleistocene bipolar temperature "seesaw" rather than CO2 increase, indicating unique limitations on glacial-age high-elevation plants may have existed due to both low temperatures and low CO2. Further development of system-specific faunal proxies can help to clarify this and other plant- and ecosystem-level responses to past environmental change.

Photosynthesis of C3, C3-C4, and C4 grasses at glacial CO2.

Most physiology comparisons of C3 and C4 plants are made under current or elevated concentrations of atmospheric CO2 which do not reflect the low CO2 environment under which C4 photosynthesis has evolved. Accordingly, photosynthetic nitrogen (PNUE) and water (PWUE) use efficiency, and the activity of the photosynthetic carboxylases [Rubisco and phosphoenolpyruvate carboxylase (PEPC)] and decarboxylases [NADP-malic enzyme (NADP-ME) and phosphoenolpyruvate carboxykinase (PEP-CK)] were compared in eight C4 grasses with NAD-ME, PCK, and NADP-ME subtypes, one C3 grass, and one C3-C4 grass grown under ambient (400 μl l(-1)) and glacial (180 μl l(-1)) CO2. Glacial CO2 caused a smaller reduction of photosynthesis and a greater increase of stomatal conductance in C4 relative to C3 and C3-C4 species. Panicum bisulcatum (C3) acclimated to glacial [CO2] by doubling Rubisco activity, while Rubisco was unchanged in Panicum milioides (C3-C4), possibly due to its high leaf N and Rubisco contents. Glacial CO2 up-regulated Rubisco and PEPC activities in concert for several C4 grasses, while NADP-ME and PEP-CK activities were unchanged, reflecting the high control exerted by the carboxylases relative to the decarboxylases on the efficiency of C4 metabolism. Despite having larger stomatal conductance at glacial CO2, C4 species maintained greater PWUE and PNUE relative to C3-C4 and C3 species due to higher photosynthetic rates. Relative to other C4 subtypes, NAD-ME and PEP-CK grasses had the highest PWUE and PNUE, respectively; relative to C3, the C3-C4 grass had higher PWUE and similar PNUE at glacial CO2. Biomass accumulation was reduced by glacial CO2 in the C3 grass relative to the C3-C4 grass, while biomass was less reduced in NAD-ME grasses compared with NADP-ME and PCK grasses. Under glacial CO2, high resource use efficiency offers a key evolutionary advantage for the transition from C3 to C4 photosynthesis in water- and nutrient-limited environments.

Carbon dioxide flux in the ablation area of Koxkar glacier, western Tien Shan, China

AbstractAccelerating loss of glacial mass caused by rising global temperatures has significant implications. The global cycle of the greenhouse gas CO2 is also associated with mineral weathering and glaciation. In glaciated areas, most estimates of atmospheric CO2 consumption are confined to chemical ionic mass balance or analog modeling methods. We applied the gradient method to the surface of Koxkar glacier, western Tien Shan, China, for a 5 month period during 2012. The overall net glacier-system CO2 exchange (NGE) rate was measured as -0.05 and -0.07mmol m–2 s–1 for regions of exposed ice and supraglacial moraine, respectively. This suggests that atmospheric CO2 drawdown may occur during ice melting because of consumption of H+ by CO2 hydrolysis that occurs in solutions. Using the degree-day model to calculate glacier ablation in bare-ice regions and considering characteristics of the NGE rate in the supraglacial debris region with the support of GIS, the daily NGE rate was estimated to be -1.23 ± 0.17μmmol m–2 d–1 between Julian days 125 and 268 of 2012. These findings present a new approach for modeling the dynamics of glacial CO2 sinks undergoing melting, and develop an understanding of the mechanism of atmospheric CO2 exchange.

Biological and climatic consequences of a cold, stratified, high latitude ocean

The flux from deep- and shallow-living radiolarian assemblages provides evidence of a glacial, high latitude, cold ocean stratification that increased biological pump efficiency and promoted ocean carbon sequestration. Greater deep (>200 m) than shallow-living (<200 m) radiolarian assemblage flux characterizes glacial North Pacific (>45° N) sediments with the deep-living Cycladophora davisiana dominant (>24%). By contrast modern radiolarian flux consists primarily of shallow-living species (C. davisiana <10%). Clues to the cause of this unusual glacial radiolarian flux come from the presently, strongly stratified Sea of Okhotsk. Here beneath a thin nutrient depleted mixed layer radiolarian and zooplankton faunas conform to the sea's physical stratification with lower concentrations of both in a Cold (−1.5 to 1 °C) Intermediate Layer (CIL) (20–125 m) and higher concentrations in waters between 200 and 500 m (Nimmergut and Abelmann, 2002). This biological stratification generates a radiolarian flux echoing that of the glacial northwest Pacific with C. davisiana 26% of total flux. Widespread C. davisiana percentages (>20%) in high latitude (>45°) glacial sediments of both hemispheres is evidence that these oceans were capped with an Okhotsk-Like Stratification (O-LS). O-LS provides mechanisms to (1) strip nutrients from surface waters depriving the deep-ocean of preformed nutrients, increasing biological pump efficiency and (2) deepen carbon re-mineralization increasing deep-ocean alkalinity. Both may have contributed to lower glacial atmospheric CO2 concentrations. O-LS would also have amplified glacial climatic cycles by promoting the spread of high latitude sea ice in winter as occurs in the Sea of Okhotsk today, and reducing gas exchange between ocean and atmosphere in summer.

Meta-analysis reveals profound responses of plant traits to glacial CO2 levels.

A general understanding of the links between atmospheric CO2 concentration and the functioning of the terrestrial biosphere requires not only an understanding of plant trait responses to the ongoing transition to higher CO2 but also the legacy effects of past low CO2. An interesting question is whether the transition from current to higher CO2 can be thought of as a continuation of the past trajectory of low to current CO2 levels. Determining this trajectory requires quantifying the effect sizes of plant response to low CO2. We performed a meta-analysis of low CO2 growth experiments on 34 studies with 54 species. We quantified how plant traits vary at reduced CO2 levels and whether C3 versus C4 and woody versus herbaceous plant species respond differently. At low CO2, plant functioning changed drastically: on average across all species, a 50% reduction in current atmospheric CO2 reduced net photosynthesis by 38%; increased stomatal conductance by 60% and decreased intrinsic water use efficiency by 48%. Total plant dry biomass decreased by 47%, while specific leaf area increased by 17%. Plant types responded similarly: the only significant differences being no increase in SLA for C4 species and a 16% smaller decrease in biomass for woody C3 species at glacial CO2. Quantitative comparison of low CO2 effect sizes to those from high CO2 studies showed that the magnitude of response of stomatal conductance, water use efficiency and SLA to increased CO2 can be thought of as continued shifts along the same line. However, net photosynthesis and dry weight responses to low CO2 were greater in magnitude than to high CO2. Understanding the causes for this discrepancy can lead to a general understanding of the links between atmospheric CO2 and plant responses with relevance for both the past and the future.

Potential role of giant marine diatoms in sequestration of atmospheric CO2 during the Last Glacial Maximum: δ13C evidence from laminated Ethmodiscus rex mats in tropical West Pacific

Giant marine diatoms, blooming or aggregating within deep chlorophyll maxima under stratified conditions, can generate substantial production and a large export flux of organic carbon from surface waters. However, their role in regulating glacial-interglacial variation in atmospheric pCO2 remains unclear. Here, we report the organic carbon isotopic compositions of Ethmodiscus rex diatoms (δ13CE. rex) and bulk sediments (δ13Corg) from a sediment core in the eastern Philippine Sea dated to ~19.5–31.0kyr B.P. and consisting of (from youngest to oldest) (1) laminated E. rex diatom mats (LDM), (2) diatomaceous clay (DC), and (3) pelagic clay (PC). Our results suggest that δ13CE. rex provides a better record of palaeoceanographic processes during LDM and DC deposition than δ13Corg because of reduced differential vital effects in near-monospecific E. rex fractions. We used the isotopic composition of the coarse E. rex fraction (δ13CE. rex (>154μm)) to calculate the CO2 partial pressure of eastern Philippine Sea surface waters (pCO2-sw) during the Last Glacial Maximum (LGM). Our pCO2-sw records suggest that the eastern Philippine Sea switched from being a strong CO2 source during DC deposition to a weak CO2 sink during LDM deposition. The role of the eastern Philippine Sea as a CO2 sink during the LGM was promoted by elevated primary production and, to a lesser extent, intensified water-column stratification. These observations highlight the potential role of giant marine diatoms in the sequestration of atmospheric CO2 during the LGM and, hence, support changes in biogenic silica fluxes as a potential cause of lower glacial atmospheric CO2. Our findings are consistent with an eolian source of silica, as proposed by the ‘silica hypothesis’ and the ‘silicon-induced alkalinity pump hypothesis’ but not by the ‘silicic acid leakage hypothesis.’

Impact of CO&amp;lt;sub&amp;gt;2&amp;lt;/sub&amp;gt; and climate on Last Glacial maximum vegetation – a factor separation

Abstract. The factor separation of Stein and Alpert (1993) is applied to simulations with the MPI Earth system model to determine the factors which cause the differences between vegetation patterns in glacial and pre-industrial climate. The factors firstly include differences in the climate, caused by a strong increase in ice masses and the radiative effect of lower greenhouse gas concentrations; secondly, differences in the ecophysiological effect of lower glacial atmospheric CO2 concentrations; and thirdly, the synergy between the pure climate effect and the pure effect of changing physiologically available CO2. It is has been shown that the synergy can be interpreted as a measure of the sensitivity of ecophysiological CO2 effect to climate. The pure climate effect mainly leads to a contraction or a shift in vegetation patterns when comparing simulated glacial and pre-industrial vegetation patterns. Raingreen shrubs benefit from the colder and drier climate. The pure ecophysiological effect of CO2 appears to be stronger than the pure climate effect for many plant functional types – in line with previous simulations. The pure ecophysiological effect of lower CO2 mainly yields a reduction in fractional coverage, a thinning of vegetation and a strong reduction in net primary production. The synergy appears to be as strong as each of the pure contributions locally, but weak on global average for most plant functional types. For tropical evergreen trees, however, the synergy is strong on global average. It diminishes the difference between glacial and pre-industrial coverage of tropical evergreen trees, due to the pure climate effect and the pure ecophysiological CO2 effect, by approximately 50 per cent.

The role of the terrestrial biosphere in the glacial CO2 problem and implilcations for future carbon-climate feedback

The role of the terrestrial biosphere in the glacial CO2 problem and implilcations for future carbon-climate feedback

Glacial CO2 cycle as a succession of key physical and biogeochemical processes

Glacial CO2 cycle as a succession of key physical and biogeochemical processes

Glacial Southern Ocean freshening at the onset of the Middle Pleistocene Climate Transition

Changes in Southern Ocean hydrography may have played an important role in the Middle Pleistocene Transition (MPT), particularly through their impact on ocean circulation and atmospheric CO2 concentrations. Here we present foraminiferal Mg/Ca and δ18O results for the subsurface dwelling planktonic foraminifer Neogloboquadrina pachyderma (sinistral) at the Ocean Drilling Program (ODP) Site 1090. Results are used to reconstruct upper ocean temperatures and derive seawater δ18O in the Subantarctic Atlantic Ocean during the MPT. The new records indicate that, starting at ∼1250ka, glacial temperatures and local (ice volume corrected) seawater δ18O in the upper water column of the Subantarctic Atlantic Ocean decreased, pointing to cooler (∼2°C) and fresher (∼0.4‰) conditions than in the preceding glacial stages. These upper ocean hydrographic changes broadly coincide with the increase in the power of the 100ky glacial–interglacial cycle in both records and with a shift towards decreased deep ventilation in the glacial Southern Ocean. Our finding suggests that an increase in Southern Ocean stratification, driven by the observed freshening of the upper water column, may have reduced the exchange of carbon between the deep Southern Ocean and the atmosphere during glacial stages. This process may have contributed, in combination with other mechanisms, to lower glacial atmospheric CO2 concentrations during the MPT.

Quantifying the ocean's role in glacial CO&amp;lt;sub&amp;gt;2&amp;lt;/sub&amp;gt; reductions

Abstract. A series of Last Glacial Maximum (LGM) marine carbon cycle sensitivity experiments is conducted to test the effect of different physical processes, as simulated by two atmosphere-ocean general circulation model (AOGCM) experiments, on atmospheric pCO2. One AOGCM solution exhibits an increase in North Atlantic Deep Water (NADW) formation under glacial conditions, whereas the other mimics an increase in Antarctic Bottom Water (AABW) associated with a weaker NADW. None of these sensitivity experiments reproduces the observed magnitude of glacial/interglacial pCO2 changes. However, to explain the reconstructed vertical gradient of dissolved inorganic carbon (DIC) of 40 mmol m−3 a marked enhancement in AABW formation is required. Furthermore, for the enhanced AABW sensitivity experiment the simulated stable carbon isotope ratio (δ13C) decreases by 0.4‰ at intermediate depths in the South Atlantic in accordance with sedimentary evidence. The shift of deep and bottom water formation sites from the North Atlantic to the Southern Ocean increases the total preformed nutrient inventory, so that the lowered efficiency of Southern Ocean nutrient utilization in turn increases atmospheric pCO2. This change eventually offsets the effect of an increased abyssal carbon pool due to stronger AABW formation. The effects of interhemispheric glacial sea-ice changes on atmospheric pCO2 oppose each other. Whereas, extended sea-ice coverage in the Southern Hemisphere reduces the air-sea gas exchange of CO2 in agreement with previous theoretical considerations, glacial advances of sea-ice in the Northern Hemisphere lead to a weakening of the oceanic carbon uptake through the physical pump. Due to enhanced gas solubility associated with lower sea surface temperature, both glacial experiments generate a reduction of atmospheric pCO2 by about 20–23 ppmv. The sensitivity experiments presented here demonstrate the presence of compensating effects of different physical processes in the ocean on glacial CO2 and the difficulty of finding a simple explanation of the glacial CO2 problem by invoking ocean dynamical changes.

Glacial CO&amp;lt;sub&amp;gt;2&amp;lt;/sub&amp;gt; cycle as a succession of key physical and biogeochemical processes

Abstract. During glacial-interglacial cycles, atmospheric CO2 concentration varied by about 100 ppmv in amplitude. While testing mechanisms that have led to the low glacial CO2 level could be done in equilibrium model experiments, an ultimate goal is to explain CO2 changes in transient simulations through the complete glacial-interglacial cycle. The computationally efficient Earth System model of intermediate complexity CLIMBER-2 is used to simulate global biogeochemistry over the last glacial cycle (126 kyr). The physical core of the model (atmosphere, ocean, land and ice sheets) is driven by orbital changes and reconstructed radiative forcing from greenhouses gases, ice, and aeolian dust. The carbon cycle model is able to reproduce the main features of the CO2 changes: a 50 ppmv CO2 drop during glacial inception, a minimum concentration at the last glacial maximum 80 ppmv lower than the Holocene value, and an abrupt 60 ppmv CO2 rise during the deglaciation. The model deep ocean δ13C also resembles reconstructions from deep-sea cores. The main drivers of atmospheric CO2 evolve in time: changes in sea surface temperatures and in the volume of bottom water of southern origin control atmospheric CO2 during the glacial inception and deglaciation; changes in carbonate chemistry and marine biology are dominant during the first and second parts of the glacial cycle, respectively. These feedback mechanisms could also significantly impact the ultimate climate response to the anthropogenic perturbation.

Onset of ‘Pacific-style’ deep-sea sedimentary carbonate cycles at the mid-Pleistocene transition

The calcium carbonate (CaCO3) content of deep-sea sediments in the Pacific Ocean increases during glacials of the late Pleistocene in comparison to interglacials, whereas records of sedimentary CaCO3 in the Atlantic Ocean show an anticorrelated pattern across glacial-interglacial cycles. Here we show that this anticorrelation in inter-ocean CaCO3 cycles arose comparatively recently, at ~1.10million years ago (Ma), during the mid-Pleistocene transition. Before this time, we show that the CaCO3 content of Pacific and Atlantic Ocean sediments experienced in-phase cyclicity, both having ‘Atlantic-style’ phasing with respect to glacial–interglacial cycles. The onset of anticorrelated cyclicity at 1.10Ma involved a twofold switch in the Pacific's CaCO3 cycles: glacial CaCO3 preservation became consistently better while interglacial preservation became consistently worse. We demonstrate that the cause of this glacial–interglacial ‘mirror imaging’ of Pacific CaCO3 dissolution cyclicity at 1.10Ma was a switching over of the relative ventilation state of abyssal South Pacific waters between glacials and interglacials. Specifically, we suggest that a consistent strengthening of deep water ventilation within the Pacific sector of the Southern Ocean during glacials drove glacial Pacific CaCO3 dissolution to diminish, while a contemporaneous weakening of very well-ventilated ‘upstream’ North Atlantic Deep Water during interglacials drove interglacial Pacific CaCO3 dissolution ‘downstream’ to intensify. We propose that the increased mean alkalinity of the global deep ocean (driven by the geographically and bathymetrically vast Pacific Ocean) during glacials after 1.10Ma may explain part of the drawdown in glacial atmospheric CO2 levels hypothesised to have been linked to the increased severity of these late Pleistocene glacials.

Atmospheric CO2 decline during the Pliocene intensification of Northern Hemisphere glaciations

Several hypotheses have been put forward to explain the onset of intensive glaciations on Greenland, Scandinavia, and North America during the Pliocene epoch between 3.6 and 2.7 million years ago (Ma). A decrease in atmospheric CO2 may have played a role during the onset of glaciations, but other tectonic and oceanic events occurring at the same time may have played a part as well. Here we present detailed atmospheric CO2 estimates from boron isotopes in planktic foraminifer shells spanning 4.6–2.0 Ma. Maximal Pliocene atmospheric CO2 estimates gradually declined from values around 410 μatm to early Pleistocene values of 300 μatm at 2.0 Ma. After the onset of large‐scale ice sheets in the Northern Hemisphere, maximal pCO2 estimates were still at 2.5 Ma +90 μatm higher than values characteristic of the early Pleistocene interglacials. By contrast, Pliocene minimal atmospheric CO2 gradually decreased from 310 to 245 μatm at 3.2 Ma, coinciding with the start of transient glaciations on Greenland. Values characteristic of early Pleistocene glacial atmospheric CO2 of 200 μatm were abruptly reached after 2.7 Ma during the late Pliocene transition. This trend is consistent with the suggestion that ocean stratification and iron fertilization increased after 2.7 Ma in the North Pacific and Southern Ocean and may have led to increased glacial CO2 storage in the oceanic abyss after 2.7 Ma onward.

Proximal trigger for late glacial Antarctic circulation and CO2 changes

Proximal trigger for late glacial Antarctic circulation and CO2 changes

Impact of CO&amp;lt;sub&amp;gt;2&amp;lt;/sub&amp;gt; and climate on the Last Glacial Maximum vegetation: results from the ORCHIDEE/IPSL models

Abstract. Vegetation reconstructions from pollen data for the Last Glacial Maximum (LGM), 21 ky ago, reveal lanscapes radically different from the modern ones, with, in particular, a massive regression of forested areas in both hemispheres. Two main factors have to be taken into account to explain these changes in comparison to today's potential vegetation: a generally cooler and drier climate and a lower level of atmospheric CO2. In order to assess the relative impact of climate and atmospheric CO2 changes on the global vegetation, we simulate the potential modern vegetation and the glacial vegetation with the dynamical global vegetation model ORCHIDEE, driven by outputs from the IPSL_CM4_v1 atmosphere-ocean general circulation model, under modern or glacial CO2 levels for photosynthesis. ORCHIDEE correctly reproduces the broad features of the glacial vegetation. Our modelling results support the view that the physiological effect of glacial CO2 is a key factor to explain vegetation changes during glacial times. In our simulations, the low atmospheric CO2 is the only driver of the tropical forests regression, and explains half of the response of temperate and boreal forests to glacial conditions. Our study shows that the sensitivity to CO2 changes depends on the background climate over a region, and also depends on the vegetation type, needleleaf trees being much more sensitive than broadleaf trees in our model. This difference of sensitivity leads to a dominance of broadleaf types in the remaining simulated forests, which is not supported by pollen data, but nonetheless suggests a potential impact of CO2 on the glacial vegetation assemblages. It also modifies the competitivity between the trees and makes the amplitude of the response to CO2 dependent on the initial vegetation state.