In Defense of Mud II: Lakes as Carbon Sinks

Mark Brenner, William F. Kenney
{"title":"In Defense of Mud II: Lakes as Carbon Sinks","authors":"Mark Brenner,&nbsp;William F. Kenney","doi":"10.1002/bes2.2187","DOIUrl":null,"url":null,"abstract":"<p>In late 1969, Edward S. Deevey, Jr. delivered a talk to the National Water Commission, with the catchy title “In Defense of Mud.” The text of his speech was published the following year in the <i>Bulletin of the Ecological Society of America</i> (Deevey <span>1970</span>). The year of publication coincided with the first celebration of Earth Day and was a time of growing realization that humans were transforming the planet, thereby threatening many species with extinction. In what may have appeared to be an odd argument for conserving non-marine aquatic ecosystems, Deevey proposed that lakes, wetlands, ponds, and estuaries deserved protection, not only because they are homes for iconic birds, mammals, and fish, but because the sediments that accumulate within them are habitats for the bacteria that reduce nitrate and sulfate, making them important players in key global biogeochemical cycles. Humans were emitting tremendous amounts of sulfate and nitrate into the atmosphere. At the same time, they were ditching and draining wetlands for agriculture and construction, thus endangering the anoxic, muddy realm in which nitrate- and sulfur-reducing microbes live. Although it is unlikely that anyone ever saw a “Save the Microbes” bumper sticker, Deevey's publication represented a novel and holistic perspective on the need to conserve continental aquatic ecosystems.</p><p>Fifty years later, Hale (<span>2020</span>) published a commentary, “In Praise of Mud,” also in the <i>Bulletin of the Ecological Society of America</i>. It expanded upon Deevey's work by providing a list of 38 reasons why humans and other animals should appreciate lake sediments, among them because they provide myriad ecosystem services, are used in the manufacture of construction and beauty products, and are rich archives of past climate and environmental information. Hale mentioned that we should also be grateful for a particular service provided by lake mud, i.e., that it “sequesters carbon, preventing its return as carbon dioxide to the atmosphere.” We agree with his claim and will focus here on that important ecosystem function. We address the issue because there exists confusion about the role lakes play in global carbon (C) cycling, particularly with respect to their being sources or sinks of C relative to the atmosphere. Given the recent rise in the concentration of CO<sub>2</sub> in the atmosphere and consequent global climate change, it is crucial that we understand the processes associated with C as it moves through inland aquatic ecosystems.</p><p>For this discussion, we use the generally accepted definition of a “sink” as it applies to any element. Broadly, an element sink can be thought of as an area of the landscape or a volume of the biosphere where the mass of an element increases over time, i.e., where inputs of the element exceed outputs. We define a C sink as a pool or compartment in the carbon biogeochemical cycle where C-containing material accumulates and is stored for a protracted period. The C sink absorbs more carbon from the atmosphere than it releases to the atmosphere. A C source, on the other hand, supplies more carbon to the atmosphere than it sequesters. We note that some engineering strategies designed to reduce C concentration in the atmosphere involve the capture of gaseous CO<sub>2</sub> and its subsequent storage, in solid form, in a subterranean sink. The percent of C stored (inputs minus outputs) relative to total C outputs has no bearing on the sink versus source determination, and it follows from our definition that the term “net sink” is redundant.</p><p>We contend that almost all lakes behave as C sinks on timescales lasting millennia, and in some cases, much longer. Lake basins continue to act as C sinks as long as they accumulate C-bearing sediment, in the form of organic matter (OM), typically ~45–50% organic C (OC) by mass, or carbonate (e.g., as CaCO<sub>3</sub>, 12% C<sub>inorg</sub> by mass), as snail, bivalve, or ostracod shells, or as photosynthetically or chemically induced precipitates. Inorganic C storage is particularly notable in lakes of karst terrains (Martin <span>2017</span>), where carbonates can account for &gt;90% of the sediment mass in shallow-water areas of waterbodies (Hodell et al. <span>2007</span>).</p><p>We felt compelled to address this topic, given the number of studies that purport to show that lakes are “sources” of C to the atmosphere. We do not dispute the fact that in many lakes, perhaps most, more gaseous CO<sub>2</sub> is transferred across the air–water interface from the water column to the atmosphere than in the opposite direction. Nonetheless, if any C that enters a lake is “permanently” stored on the lake bottom, in any form, we argue that the basin functions as a C sink. That is, the lake is a C sink if there is a growing amount of C being accumulated and stored below the area circumscribed by the lake shoreline.</p><p>There are several possible reasons why some investigators have concluded that lakes are not C sinks: (1) they misunderstand the definition of the term “sink” or adopt a modified definition, and/or (2) they use inappropriate measures to determine if a lake is accumulating C. For example (a) they look at partial pressure of CO<sub>2</sub> in surface waters, or pH and alkalinity values, simply to evaluate whether dissolved CO<sub>2</sub> in the water column will tend to evolve to the atmosphere, (b) they directly measure gas exchange across the air-water interface, or (c) they look at the ratio of CO<sub>2</sub> uptake during lacustrine primary production (photosynthesis) to carbon dioxide gas produced by community respiration in the water column (P/R). In such studies, accruing C-containing sediments are typically ignored. Lastly, investigators may decide that some arbitrary proportion of C fixed in the lake must be sequestered in the sediments to rightfully qualify the waterbody as a C sink. This requirement resulted in the contradictory claim that subtropical stormwater ponds are net sources of C to the atmosphere at the same time that they continue to accumulate ever greater amounts of C through time (Goeckner et al. <span>2022</span>). In contrast, a study of temperate artificial ponds determined that those waterbodies displayed high OC sequestration rates, and suggested that global OC burial in inland waters is probably underestimated (Holgerson et al. <span>2024</span>).</p><p>We are not the first to address the confusion about the role of lakes as “sources vs. sinks” in the global C cycle. Dean and Gorham (<span>1998</span>) mentioned that some studies had suggested lakes were sources of CO<sub>2</sub> in the atmosphere (Cole et al. <span>1994</span>, Molot and Dillon <span>1997</span>), but they then went on to make the case that sedimentation in inland waterbodies stored substantial C and that lakes and wetlands were thus C sinks. The authors estimated that lakes, reservoirs, and peatlands around the world accumulate OC at a rate of ~300 Tg/year, whereas OC accumulation in the oceans amounts to ~100 Tg/year. By their reckoning, lakes account for ~42 Tg/year (14%) of the total OC stored annually in inland lentic aquatic ecosystems.</p><p>Much of the recent interest in C cycling in lakes is related to how climate and human activities influence C storage in (Anderson et al. <span>2020</span>) or loss from lakes (St. Louis et al. <span>2000</span>, Kosten et al. <span>2010</span>). It has been noted that the C cycle, as it applies to lakes, both regulates and is regulated by climate change (Tranvik et al. <span>2009</span>, Yasarer <span>2015</span>). Carbon enters lakes in gaseous form across the air-water interface, as dissolved inorganic C (DIC) in rainfall and other input waters, and as dissolved or particulate organic C (DOC/POC) from the watershed and airshed. Carbon that is fixed via photosynthesis or bio-precipitated within the lake ultimately came from outside the waterbody (i.e., from the atmosphere). It is indisputable that large amounts of gaseous C are lost from lakes to the atmosphere, in the form of CO<sub>2</sub> and methane (CH<sub>4</sub>). Some lakes are even thought to be heterotrophic (Bachmann et al. <span>2000</span>), i.e., they display higher rates of water column respiration (CO<sub>2</sub> generation) than primary production (CO<sub>2</sub> consumption). We note that even such heterotrophic lakes can function as C sinks, as long as they continue to accumulate carbonaceous sediment. Indeed, most lakes probably evolve almost all the C that enters their water column back to the atmosphere, whether the C in the water column is present as DIC, allochthonous DOC/POC, or C fixed within the lake by algae and macrophytes, the latter representing POC that was fixed by photosynthesis, using lake DIC that ultimately came from the atmosphere. It is important, however, to keep in mind that the C in the lake, whether allochthonous DIC, DOC, POC, or fixed in situ via photosynthesis (autochthonous), ultimately came from the atmosphere. As some have suggested, it might be more appropriate to view lakes as “conduits” through which substantial C passes on its way back to the atmosphere (Tranvik et al. <span>2009</span>, Yasarer <span>2015</span>).</p><p>The case for lakes being C sinks can benefit from drawing an analogy to household economies (Fig. 1A, B). Although an imperfect comparison, it may be helpful to think of the money that enters and leaves household accounts as analogous to C flow into and out of lakes. Just as lakes receive C from multiple external sources, household income can come from wages, interest, investments, etc. Some hobbyists may sell artisanal wares like ceramics and jewelry that they fabricate at home, which can be thought of as an autochthonous (in-house) revenue source. As is the case for C loss from lakes, a large proportion of household income leaves home accounts to cover living expenses, i.e., necessities like rent or mortgage payments, groceries, clothing, furniture, utilities, car payments, etc., as well as discretionary expenditures such as vacations and entertainment. A wise money manager will, however, arrange for some portion of their income to be earmarked for use later in life. That money can be put into savings or retirement accounts, stocks and bonds, etc., all of which, one hopes, will continue to grow in value over time, i.e., accrue more money. Depending on monthly household income and financial obligations, smaller or larger sums of money can be earmarked for savings. Money deposited in savings is temporarily (decades?) removed from circulation in the economy, just as C in lake-bottom sediments is prevented from cycling back into the atmosphere, often on timescales lasting thousands of years.</p><p>The household economy analogy may be imperfect for several reasons, perhaps most obviously because human life spans and the life spans of lakes on the landscape are so different. After leaving their jobs, retirees may begin to draw down their savings accounts, which they rely on when they no longer receive salary, rendering those accounts a money source to the economy rather than a sink. To further analogize, we can envision circumstances in which accumulated carbonaceous lake sediments cease to accrue and then become sources of C to the atmosphere: (1) shallow (ephemeral) lakes may dry completely, exposing their accumulated organic and carbonate deposits to sub-aerial conditions and oxidation/dissolution, (2) accumulated peat deposits may be mined and combusted for fuel or used as soil amendments, (3) wetland soils may be ditched and drained for cropping and animal grazing, and (4) lake sediments may be dredged to enhance navigation or improve water quality. But these examples are exceptions to the general rule. Almost all lakes around the world, most of which formed since the last deglaciation, continue to hold water and accumulate C-bearing sediments.</p><p>Perhaps one way to address the “source/sink conundrum” is to pose the following hypothetical question: If a lake were to sequester in its sediments all the C that entered its water column, i.e., if it were 100% efficient at storing incoming C on the lake bottom, could we all agree that the lake behaves as a C sink? We suspect that most or all individuals would respond in the affirmative. We can now pose the same question for a lake that is 75% efficient at storing C in its sediments, i.e., it permanently buries three-quarters of all the incoming C. Again, most respondents would probably agree that such a lake is a C sink. If we reduce the efficiency to 50%, half of all incoming C is preserved on the lake bottom, and half makes its way back to the atmosphere. Here we might refer to our household economy analogy and ask, who would not be happy to be able to put half of their annual income into a savings account? Let's now entertain the same question for lakes that sequester in their sediment just a few percent of their incoming C (analogous to the mean percent of disposable personal income that US residents put into savings, ~4%). This is probably the case for most lakes, and we contend that they are nevertheless C sinks; places where C is stored on timescales of years to many millennia. We also argue that the assignment of an arbitrary criterion, in terms of percent of incoming C stored, cannot be used to determine if a lake is a C source or sink.</p><p>We began this discussion by presenting our definition of a C sink to avoid semantic ambiguity. It is our contention that the “source/sink debate” as applied to inland waterbodies, diverts attention from interesting questions about the role of lakes in C cycling (Cole et al. <span>2007</span>). Fortunately, many who study continental aquatic ecosystems, especially paleolimnologists, i.e., scientists who focus on the sediment compartment where C is stored, are addressing those questions. To list just a few such questions: (1) how do C accumulation rates differ spatially within lakes? (Lin et al. <span>2022</span>), (2) how do C accumulation rates differ among lakes, with respect to trophic state, temperature, and dominant primary producer community (macrophytes versus phytoplankton)? (Kosten et al. <span>2010</span>, Balmer and Downing <span>2011</span>, Alcocer et al. <span>2014</span>, Heathcote et al. <span>2015</span>, Reed et al. <span>2018</span>, Waters et al. <span>2019</span>, Anderson et al. <span>2020</span>), (3) how has C sequestration in lakes changed throughout the Holocene? (Kenney et al. <span>2016</span>), (4) how does C storage on lake bottoms change in response to land clearance? (Heathcote and Downing <span>2012</span>, Dietz et al. <span>2015</span>), (5) what are the principal sources (allochthonous/autochthonous) of C in lake sediments? (Meyers and Ishiwatari <span>1993</span>), (6) what factors affect the efficiency of C sequestration [storage/input] in lakes? (Tranvik et al. <span>2009</span>), (7) what is the magnitude of CH<sub>4</sub> evolution from lakes? (Johnson et al. <span>2022</span>), (8) what is the importance of lake and reservoir C storage in the global C biogeochemical cycle (Mendonça et al. <span>2017</span>)? and how do terrestrial and aquatic biogeochemical processes in karst terrains influence C flux to and from the atmosphere (Chen et al. <span>2023</span>)? As we continue to further address those questions, it will be helpful to keep in mind that lake sediments are C sinks and that continental water bodies play an important role in sequestering C and keeping it out of the atmosphere.</p>","PeriodicalId":93418,"journal":{"name":"Bulletin of the Ecological Society of America","volume":"106 1","pages":""},"PeriodicalIF":0.0000,"publicationDate":"2024-10-08","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://onlinelibrary.wiley.com/doi/epdf/10.1002/bes2.2187","citationCount":"0","resultStr":null,"platform":"Semanticscholar","paperid":null,"PeriodicalName":"Bulletin of the Ecological Society of America","FirstCategoryId":"1085","ListUrlMain":"https://onlinelibrary.wiley.com/doi/10.1002/bes2.2187","RegionNum":0,"RegionCategory":null,"ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":null,"EPubDate":"","PubModel":"","JCR":"","JCRName":"","Score":null,"Total":0}
引用次数: 0

Abstract

In late 1969, Edward S. Deevey, Jr. delivered a talk to the National Water Commission, with the catchy title “In Defense of Mud.” The text of his speech was published the following year in the Bulletin of the Ecological Society of America (Deevey 1970). The year of publication coincided with the first celebration of Earth Day and was a time of growing realization that humans were transforming the planet, thereby threatening many species with extinction. In what may have appeared to be an odd argument for conserving non-marine aquatic ecosystems, Deevey proposed that lakes, wetlands, ponds, and estuaries deserved protection, not only because they are homes for iconic birds, mammals, and fish, but because the sediments that accumulate within them are habitats for the bacteria that reduce nitrate and sulfate, making them important players in key global biogeochemical cycles. Humans were emitting tremendous amounts of sulfate and nitrate into the atmosphere. At the same time, they were ditching and draining wetlands for agriculture and construction, thus endangering the anoxic, muddy realm in which nitrate- and sulfur-reducing microbes live. Although it is unlikely that anyone ever saw a “Save the Microbes” bumper sticker, Deevey's publication represented a novel and holistic perspective on the need to conserve continental aquatic ecosystems.

Fifty years later, Hale (2020) published a commentary, “In Praise of Mud,” also in the Bulletin of the Ecological Society of America. It expanded upon Deevey's work by providing a list of 38 reasons why humans and other animals should appreciate lake sediments, among them because they provide myriad ecosystem services, are used in the manufacture of construction and beauty products, and are rich archives of past climate and environmental information. Hale mentioned that we should also be grateful for a particular service provided by lake mud, i.e., that it “sequesters carbon, preventing its return as carbon dioxide to the atmosphere.” We agree with his claim and will focus here on that important ecosystem function. We address the issue because there exists confusion about the role lakes play in global carbon (C) cycling, particularly with respect to their being sources or sinks of C relative to the atmosphere. Given the recent rise in the concentration of CO2 in the atmosphere and consequent global climate change, it is crucial that we understand the processes associated with C as it moves through inland aquatic ecosystems.

For this discussion, we use the generally accepted definition of a “sink” as it applies to any element. Broadly, an element sink can be thought of as an area of the landscape or a volume of the biosphere where the mass of an element increases over time, i.e., where inputs of the element exceed outputs. We define a C sink as a pool or compartment in the carbon biogeochemical cycle where C-containing material accumulates and is stored for a protracted period. The C sink absorbs more carbon from the atmosphere than it releases to the atmosphere. A C source, on the other hand, supplies more carbon to the atmosphere than it sequesters. We note that some engineering strategies designed to reduce C concentration in the atmosphere involve the capture of gaseous CO2 and its subsequent storage, in solid form, in a subterranean sink. The percent of C stored (inputs minus outputs) relative to total C outputs has no bearing on the sink versus source determination, and it follows from our definition that the term “net sink” is redundant.

We contend that almost all lakes behave as C sinks on timescales lasting millennia, and in some cases, much longer. Lake basins continue to act as C sinks as long as they accumulate C-bearing sediment, in the form of organic matter (OM), typically ~45–50% organic C (OC) by mass, or carbonate (e.g., as CaCO3, 12% Cinorg by mass), as snail, bivalve, or ostracod shells, or as photosynthetically or chemically induced precipitates. Inorganic C storage is particularly notable in lakes of karst terrains (Martin 2017), where carbonates can account for >90% of the sediment mass in shallow-water areas of waterbodies (Hodell et al. 2007).

We felt compelled to address this topic, given the number of studies that purport to show that lakes are “sources” of C to the atmosphere. We do not dispute the fact that in many lakes, perhaps most, more gaseous CO2 is transferred across the air–water interface from the water column to the atmosphere than in the opposite direction. Nonetheless, if any C that enters a lake is “permanently” stored on the lake bottom, in any form, we argue that the basin functions as a C sink. That is, the lake is a C sink if there is a growing amount of C being accumulated and stored below the area circumscribed by the lake shoreline.

There are several possible reasons why some investigators have concluded that lakes are not C sinks: (1) they misunderstand the definition of the term “sink” or adopt a modified definition, and/or (2) they use inappropriate measures to determine if a lake is accumulating C. For example (a) they look at partial pressure of CO2 in surface waters, or pH and alkalinity values, simply to evaluate whether dissolved CO2 in the water column will tend to evolve to the atmosphere, (b) they directly measure gas exchange across the air-water interface, or (c) they look at the ratio of CO2 uptake during lacustrine primary production (photosynthesis) to carbon dioxide gas produced by community respiration in the water column (P/R). In such studies, accruing C-containing sediments are typically ignored. Lastly, investigators may decide that some arbitrary proportion of C fixed in the lake must be sequestered in the sediments to rightfully qualify the waterbody as a C sink. This requirement resulted in the contradictory claim that subtropical stormwater ponds are net sources of C to the atmosphere at the same time that they continue to accumulate ever greater amounts of C through time (Goeckner et al. 2022). In contrast, a study of temperate artificial ponds determined that those waterbodies displayed high OC sequestration rates, and suggested that global OC burial in inland waters is probably underestimated (Holgerson et al. 2024).

We are not the first to address the confusion about the role of lakes as “sources vs. sinks” in the global C cycle. Dean and Gorham (1998) mentioned that some studies had suggested lakes were sources of CO2 in the atmosphere (Cole et al. 1994, Molot and Dillon 1997), but they then went on to make the case that sedimentation in inland waterbodies stored substantial C and that lakes and wetlands were thus C sinks. The authors estimated that lakes, reservoirs, and peatlands around the world accumulate OC at a rate of ~300 Tg/year, whereas OC accumulation in the oceans amounts to ~100 Tg/year. By their reckoning, lakes account for ~42 Tg/year (14%) of the total OC stored annually in inland lentic aquatic ecosystems.

Much of the recent interest in C cycling in lakes is related to how climate and human activities influence C storage in (Anderson et al. 2020) or loss from lakes (St. Louis et al. 2000, Kosten et al. 2010). It has been noted that the C cycle, as it applies to lakes, both regulates and is regulated by climate change (Tranvik et al. 2009, Yasarer 2015). Carbon enters lakes in gaseous form across the air-water interface, as dissolved inorganic C (DIC) in rainfall and other input waters, and as dissolved or particulate organic C (DOC/POC) from the watershed and airshed. Carbon that is fixed via photosynthesis or bio-precipitated within the lake ultimately came from outside the waterbody (i.e., from the atmosphere). It is indisputable that large amounts of gaseous C are lost from lakes to the atmosphere, in the form of CO2 and methane (CH4). Some lakes are even thought to be heterotrophic (Bachmann et al. 2000), i.e., they display higher rates of water column respiration (CO2 generation) than primary production (CO2 consumption). We note that even such heterotrophic lakes can function as C sinks, as long as they continue to accumulate carbonaceous sediment. Indeed, most lakes probably evolve almost all the C that enters their water column back to the atmosphere, whether the C in the water column is present as DIC, allochthonous DOC/POC, or C fixed within the lake by algae and macrophytes, the latter representing POC that was fixed by photosynthesis, using lake DIC that ultimately came from the atmosphere. It is important, however, to keep in mind that the C in the lake, whether allochthonous DIC, DOC, POC, or fixed in situ via photosynthesis (autochthonous), ultimately came from the atmosphere. As some have suggested, it might be more appropriate to view lakes as “conduits” through which substantial C passes on its way back to the atmosphere (Tranvik et al. 2009, Yasarer 2015).

The case for lakes being C sinks can benefit from drawing an analogy to household economies (Fig. 1A, B). Although an imperfect comparison, it may be helpful to think of the money that enters and leaves household accounts as analogous to C flow into and out of lakes. Just as lakes receive C from multiple external sources, household income can come from wages, interest, investments, etc. Some hobbyists may sell artisanal wares like ceramics and jewelry that they fabricate at home, which can be thought of as an autochthonous (in-house) revenue source. As is the case for C loss from lakes, a large proportion of household income leaves home accounts to cover living expenses, i.e., necessities like rent or mortgage payments, groceries, clothing, furniture, utilities, car payments, etc., as well as discretionary expenditures such as vacations and entertainment. A wise money manager will, however, arrange for some portion of their income to be earmarked for use later in life. That money can be put into savings or retirement accounts, stocks and bonds, etc., all of which, one hopes, will continue to grow in value over time, i.e., accrue more money. Depending on monthly household income and financial obligations, smaller or larger sums of money can be earmarked for savings. Money deposited in savings is temporarily (decades?) removed from circulation in the economy, just as C in lake-bottom sediments is prevented from cycling back into the atmosphere, often on timescales lasting thousands of years.

The household economy analogy may be imperfect for several reasons, perhaps most obviously because human life spans and the life spans of lakes on the landscape are so different. After leaving their jobs, retirees may begin to draw down their savings accounts, which they rely on when they no longer receive salary, rendering those accounts a money source to the economy rather than a sink. To further analogize, we can envision circumstances in which accumulated carbonaceous lake sediments cease to accrue and then become sources of C to the atmosphere: (1) shallow (ephemeral) lakes may dry completely, exposing their accumulated organic and carbonate deposits to sub-aerial conditions and oxidation/dissolution, (2) accumulated peat deposits may be mined and combusted for fuel or used as soil amendments, (3) wetland soils may be ditched and drained for cropping and animal grazing, and (4) lake sediments may be dredged to enhance navigation or improve water quality. But these examples are exceptions to the general rule. Almost all lakes around the world, most of which formed since the last deglaciation, continue to hold water and accumulate C-bearing sediments.

Perhaps one way to address the “source/sink conundrum” is to pose the following hypothetical question: If a lake were to sequester in its sediments all the C that entered its water column, i.e., if it were 100% efficient at storing incoming C on the lake bottom, could we all agree that the lake behaves as a C sink? We suspect that most or all individuals would respond in the affirmative. We can now pose the same question for a lake that is 75% efficient at storing C in its sediments, i.e., it permanently buries three-quarters of all the incoming C. Again, most respondents would probably agree that such a lake is a C sink. If we reduce the efficiency to 50%, half of all incoming C is preserved on the lake bottom, and half makes its way back to the atmosphere. Here we might refer to our household economy analogy and ask, who would not be happy to be able to put half of their annual income into a savings account? Let's now entertain the same question for lakes that sequester in their sediment just a few percent of their incoming C (analogous to the mean percent of disposable personal income that US residents put into savings, ~4%). This is probably the case for most lakes, and we contend that they are nevertheless C sinks; places where C is stored on timescales of years to many millennia. We also argue that the assignment of an arbitrary criterion, in terms of percent of incoming C stored, cannot be used to determine if a lake is a C source or sink.

We began this discussion by presenting our definition of a C sink to avoid semantic ambiguity. It is our contention that the “source/sink debate” as applied to inland waterbodies, diverts attention from interesting questions about the role of lakes in C cycling (Cole et al. 2007). Fortunately, many who study continental aquatic ecosystems, especially paleolimnologists, i.e., scientists who focus on the sediment compartment where C is stored, are addressing those questions. To list just a few such questions: (1) how do C accumulation rates differ spatially within lakes? (Lin et al. 2022), (2) how do C accumulation rates differ among lakes, with respect to trophic state, temperature, and dominant primary producer community (macrophytes versus phytoplankton)? (Kosten et al. 2010, Balmer and Downing 2011, Alcocer et al. 2014, Heathcote et al. 2015, Reed et al. 2018, Waters et al. 2019, Anderson et al. 2020), (3) how has C sequestration in lakes changed throughout the Holocene? (Kenney et al. 2016), (4) how does C storage on lake bottoms change in response to land clearance? (Heathcote and Downing 2012, Dietz et al. 2015), (5) what are the principal sources (allochthonous/autochthonous) of C in lake sediments? (Meyers and Ishiwatari 1993), (6) what factors affect the efficiency of C sequestration [storage/input] in lakes? (Tranvik et al. 2009), (7) what is the magnitude of CH4 evolution from lakes? (Johnson et al. 2022), (8) what is the importance of lake and reservoir C storage in the global C biogeochemical cycle (Mendonça et al. 2017)? and how do terrestrial and aquatic biogeochemical processes in karst terrains influence C flux to and from the atmosphere (Chen et al. 2023)? As we continue to further address those questions, it will be helpful to keep in mind that lake sediments are C sinks and that continental water bodies play an important role in sequestering C and keeping it out of the atmosphere.

Abstract Image

为泥浆辩护2:湖泊作为碳汇
1969年底,小爱德华·s·迪维(Edward S. Deevey, Jr.)在国家水资源委员会(National Water Commission)做了一次演讲,题目很吸引人:“为泥浆辩护”(In Defense of Mud)。次年,他的演讲全文发表在《美国生态学会公报》(Deevey 1970)上。这本书出版的那一年恰逢第一次庆祝地球日,人们越来越意识到人类正在改变地球,从而威胁到许多物种的灭绝。对于保护非海洋水生生态系统,Deevey提出湖泊、湿地、池塘和河口值得保护,不仅因为它们是标志性鸟类、哺乳动物和鱼类的家园,而且因为其中积累的沉积物是细菌的栖息地,可以减少硝酸盐和硫酸盐,使它们在关键的全球生物地球化学循环中发挥重要作用。人类向大气中排放了大量的硫酸盐和硝酸盐。与此同时,为了农业和建筑,他们在湿地开凿沟渠,排干水,从而危及了硝酸盐和硫还原微生物赖以生存的缺氧、泥泞的环境。虽然不太可能有人见过“拯救微生物”的汽车保险杠贴纸,但迪维的出版物代表了一种新颖而全面的观点,即保护大陆水生生态系统的必要性。五十年后,黑尔(2020)在《美国生态学会公报》上发表了一篇评论文章《赞美泥浆》。它扩展了Deevey的工作,提供了38个人类和其他动物应该欣赏湖泊沉积物的理由,其中包括它们提供了无数的生态系统服务,用于制造建筑和美容产品,并且是过去气候和环境信息的丰富档案。黑尔提到,我们还应该感谢湖泥提供的一项特殊服务,即它“隔离碳,防止其以二氧化碳的形式返回大气”。我们同意他的观点,并将在此重点讨论重要的生态系统功能。我们之所以讨论这个问题,是因为人们对湖泊在全球碳(C)循环中所起的作用存在困惑,特别是关于它们相对于大气是碳的来源还是汇。鉴于最近大气中二氧化碳浓度的上升以及随之而来的全球气候变化,我们必须了解碳在内陆水生生态系统中移动的相关过程。在本讨论中,我们使用普遍接受的“接收器”定义,因为它适用于任何元素。从广义上讲,元素汇可以被认为是景观的一个区域或生物圈的一个体积,其中元素的质量随着时间的推移而增加,即元素的输入超过输出。我们将碳汇定义为碳生物地球化学循环中的一个池或隔间,含碳物质在其中积累并长期储存。碳汇从大气中吸收的碳比释放到大气中的碳多。另一方面,碳源向大气提供的碳比它吸收的碳多。我们注意到,一些旨在降低大气中碳浓度的工程策略涉及捕获气态二氧化碳并随后将其以固体形式储存在地下汇中。存储的C(输入减去输出)相对于总C输出的百分比与汇与源的确定无关,并且根据我们的定义,术语“净汇”是多余的。我们认为,几乎所有的湖泊都像碳汇一样,在数千年的时间尺度上,在某些情况下,时间要长得多。湖盆继续作为碳汇,只要它们积累含碳沉积物,以有机质(OM)的形式,通常为45-50%有机碳(OC)的质量,或碳酸盐(如CaCO3, 12%的质量),蜗牛壳,双壳类,或介形虫壳,或光合作用或化学诱导的沉淀。无机碳储存在喀斯特地形的湖泊中尤为显著(Martin 2017),其中碳酸盐可以占水体浅水区沉积物质量的90% (Hodell et al. 2007)。考虑到大量的研究表明湖泊是大气中碳的“来源”,我们觉得有必要讨论这个话题。在许多湖泊中,也许是大多数湖泊中,通过空气-水界面从水柱向大气输送的气态二氧化碳比从水柱向大气输送的二氧化碳要多,这一事实是毋庸置疑的。尽管如此,如果任何进入湖泊的碳以任何形式“永久”储存在湖底,我们认为盆地的功能是碳汇。也就是说,如果在湖岸线划定的区域以下有越来越多的碳积累和储存,那么湖泊就是一个碳汇。 一些研究人员得出湖泊不是碳汇的结论,可能有以下几个原因:(1)他们误解了“汇”一词的定义或采用了修改后的定义,和/或(2)他们使用了不适当的措施来确定湖泊是否正在积聚c。例如(a)他们观察地表水中CO2的分压,或pH值和碱度值,仅仅是为了评估水柱中溶解的CO2是否倾向于进化到大气中,(b)他们直接测量空气-水界面上的气体交换。或者(c)他们观察湖泊初级生产(光合作用)期间二氧化碳吸收与水体中群落呼吸产生的二氧化碳气体的比率(P/R)。在这类研究中,积累的含碳沉积物通常被忽略。最后,调查人员可能会认为,固定在湖中的任意比例的碳必须被隔离在沉积物中,从而使水体有资格成为碳汇。这一要求导致了一种相互矛盾的说法,即亚热带雨水池是大气中碳的净来源,同时随着时间的推移,它们继续积累越来越多的碳(Goeckner et al. 2022)。相比之下,一项对温带人工池塘的研究确定,这些水体显示出较高的碳固存率,并表明全球内陆水域的碳埋藏可能被低估了(Holgerson et al. 2024)。我们并不是第一个解决湖泊在全球C循环中作为“源与汇”角色的困惑的人。Dean和Gorham(1998)提到,一些研究表明湖泊是大气中二氧化碳的来源(Cole et al. 1994, Molot and Dillon 1997),但他们接着提出,内陆水体的沉积储存了大量的碳,因此湖泊和湿地是碳汇。作者估计,世界各地的湖泊、水库和泥炭地以每年~300 Tg的速度积累OC,而海洋中的OC积累量为每年~100 Tg。根据他们的估算,湖泊占内陆水生生态系统年总OC储存量的约42 Tg/年(14%)。最近对湖泊中碳循环的兴趣主要与气候和人类活动如何影响湖泊中的碳储存(Anderson et al. 2020)或湖泊中的碳损失(St. Louis et al. 2000, Kosten et al. 2010)有关。已经注意到,C循环,因为它适用于湖泊,既调节又受气候变化的调节(Tranvik et al. 2009, Yasarer 2015)。碳以气态形式通过空气-水界面进入湖泊,作为降雨和其他输入水中溶解的无机碳(DIC),以及作为流域和大气中的溶解或颗粒有机碳(DOC/POC)。通过光合作用或湖泊内生物沉淀固定的碳最终来自水体外部(即来自大气)。无可争辩的是,大量的气态碳以二氧化碳和甲烷(CH4)的形式从湖泊散失到大气中。有些湖泊甚至被认为是异养的(Bachmann et al. 2000),即它们的水柱呼吸(CO2产生)速率高于初级生产(CO2消耗)速率。我们注意到,即使是这样的异养湖泊也可以作为碳汇,只要它们继续积累碳质沉积物。事实上,大多数湖泊可能会将进入水柱的几乎所有碳都进化回大气,无论水柱中的碳是作为DIC、外来DOC/POC存在,还是由藻类和大型植物固定在湖中的C,后者代表通过光合作用固定的POC,利用最终来自大气的湖泊DIC。然而,重要的是要记住,湖泊中的C,无论是外来的DIC、DOC、POC,还是通过光合作用原位固定的(本地的),最终都来自大气。正如一些人所建议的那样,将湖泊视为“管道”可能更合适,大量碳通过这些管道返回大气(Tranvik等人2009,Yasarer 2015)。湖泊是碳汇的情况可以从与家庭经济的类比中受益(图1A, B)。尽管这是一个不完美的比较,但将进入和离开家庭账户的钱比作流入和流出湖泊的碳流可能会有所帮助。就像湖泊从多种外部来源获得C一样,家庭收入可以来自工资、利息、投资等。一些爱好者可能会出售他们在家里制作的陶瓷和珠宝等手工制品,这可以被认为是一种本地(内部)收入来源。与湖泊C损失的情况一样,家庭收入的很大一部分用于支付生活费用,即租金或抵押贷款、杂货、服装、家具、水电费、汽车付款等必需品,以及度假和娱乐等可自由支配的支出。 然而,明智的理财经理会安排一部分收入专门用于以后的生活。这些钱可以存入储蓄或退休账户、股票和债券等,人们希望所有这些都能随着时间的推移继续增值,也就是说,积累更多的钱。根据每月的家庭收入和财政负担,或多或少的钱可以被指定为储蓄。存放在储蓄中的钱会暂时(几十年?)从经济循环中移走,就像湖底沉积物中的碳被阻止循环回到大气中一样,通常是在数千年的时间尺度上。家庭经济的类比可能有几个不完美的原因,也许最明显的原因是人类的寿命和湖泊的寿命在景观上是如此不同。离开工作岗位后,退休人员可能会开始提取他们的储蓄账户,当他们不再领取工资时,他们就会依赖这些账户,使这些账户成为经济的资金来源,而不是一个水池。为了进一步类比,我们可以设想这样一种情况:积累的含碳湖泊沉积物停止积累,然后成为大气中碳的来源:(1)浅湖(短暂)可能完全干涸,使其积累的有机和碳酸盐沉积物暴露于地面条件和氧化/溶解;(2)积累的泥炭沉积物可能被开采和燃烧为燃料或用作土壤改良剂;(3)湿地土壤可能被挖沟和排干,用于种植和动物放牧;(4)湖泊沉积物可能被疏浚,以加强航行或改善水质。但这些例子都是一般规律的例外。世界上几乎所有的湖泊,其中大部分是在最后一次冰川消融后形成的,仍然保持着水并积累着含碳沉积物。也许解决“源/汇难题”的一种方法是提出以下假设问题:如果一个湖泊将进入其水柱的所有碳都封存在沉积物中,也就是说,如果它100%有效地将进入的碳储存在湖底,我们是否都同意湖泊的行为是一个碳汇?我们猜想,大多数或所有人都会作出肯定的答复。我们现在可以对一个在沉积物中储存C的效率为75%的湖泊提出同样的问题,也就是说,它永久地埋葬了所有进入的C的四分之三。再次,大多数受访者可能会同意这样的湖泊是一个C汇。如果我们将效率降低到50%,那么所有进入的碳中有一半被保存在湖底,另一半会回到大气中。在这里,我们可以参考我们的家庭经济类比并问,谁会不高兴能够将他们年收入的一半存入储蓄账户?现在让我们考虑一下同样的问题,对于那些在沉积物中只吸收了流入碳的百分之几的湖泊(类似于美国居民储蓄的可支配个人收入的平均百分比,约4%)。大多数湖泊的情况可能都是这样,我们认为它们仍然是碳汇;在这些地方,碳的储存时间从几年到几千年不等。我们还认为,一个任意标准的分配,以输入的C存储的百分比,不能用来确定一个湖是C源还是C汇。为了避免语义歧义,我们首先给出了C汇的定义。我们的论点是,“源/汇之争”适用于内陆水体,转移了人们对湖泊在碳循环中作用的有趣问题的注意力(Cole et al. 2007)。幸运的是,许多研究大陆水生生态系统的人,特别是古湖泊学家,即专注于储存碳的沉积物隔间的科学家,正在解决这些问题。举几个这样的问题:(1)湖泊内碳积累速率的空间差异是怎样的?(Lin et al. 2022),(2)在营养状态、温度和主要初级生产者群落(大型植物与浮游植物)方面,湖泊之间的碳积累速率有何差异?(Kosten等人,2010年,Balmer和Downing 2011年,Alcocer等人,2014年,Heathcote等人,2015年,Reed等人,2018年,Waters等人,2019年,Anderson等人,2020年),(3)整个全新世湖泊中的碳封存是如何变化的?(Kenney et al. 2016),(4)湖底碳储量如何随土地清理而变化?(Heathcote and Downing 2012, Dietz et al. 2015),(5)湖泊沉积物中C的主要来源(外来/本地)是什么?(Meyers and Ishiwatari 1993),(6)影响湖泊碳固存[储存/输入]效率的因素是什么?(Tranvik et al. 2009),(7)湖泊CH4演化的幅度是多少?(Johnson et al. 2022),(8)湖泊和水库碳储量在全球碳生物地球化学循环中的重要性是什么(mendonpada et al. 2017)?以及喀斯特地形的陆地和水生生物地球化学过程如何影响进出大气的碳通量(Chen等)。 2023) ?当我们继续进一步解决这些问题时,记住湖泊沉积物是碳汇,而大陆水体在隔离碳并将其排除在大气之外方面发挥着重要作用,将会有所帮助。
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