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Dissolved Organic Matter in Oceanic Waters

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Dissolved Organic Matter in Oceanic Waters Journal of Oceanography, Vol. 59, pp. 129 to 147, 2003 Review Dissolved Organic Matter in Oceanic Waters 1 2 HIROSHI OGAWA * and EIICHIRO TANOUE 1Ocean Research Institute, University of Tokyo, Minamidai, Nakano-ku, Tokyo 164-8639, Japan 2Graduate School of Environmental Studies, Nagoya University, Furo-cho, Chigusa-ku, Nagoya 464-8601, Japan (Received 12 February 2002; in revised form 3 July 2002; accepted 3 July 2002) The amount of information on oceanic dissolved organic matter (DOM) has increased Keywords: dramatically in the last decade thanks to the advances in chemical characterization. ⋅ Dissolved organic This information has supported the development of some novel and important ideas matter (DOM), ⋅ for DOM dynamics in the ocean. Consequently, we have a better understanding of dissolved organic the importance of DOM in oceanic biogeochemical cycles. Here we review studies carbon (DOC), ⋅ carbon cycle, published mainly during 1995–2001, synthesize them and discuss unsolved problems ⋅ molecular weight and future challenges. The measurement, distribution and turnover of dissolved or- distribution, ganic carbon (DOC) are presented as the bulk dynamics of the oceanic DOM. The ⋅ C:N ratio, size spectrum, elemental composition, and chemical compositions at molecular and ⋅ chemical composi- functional group levels are described. The mechanisms proposed for the survival of tion, biomolecules in DOM are discussed. ⋅ biomolecules. 1. Introduction organic carbon occurs as non-living DOC (e.g., Cauwet, Dissolved organic matter (DOM) in the sea is one of 1979). In contrast to the organic reservoirs on land, the the largest reservoirs of organic matter on the earth’s sur- processes by which DOM has been formed are unclear, face (others include soil organic matter and plant biomass and actual sources and the chemical nature of DOM are on land), holding approximately as much carbon as is not well known. available in atmospheric carbon dioxide (Hedges, 1992). DOM is still the least understood organic reservoir The fact that DOM is a huge organic reservoir on the on the earth, but our knowledge of DOM has been rap- earth’s surface has continued to impel ocean scientists to idly increasing. Recent advancements in DOM study have investigate what DOM is, in terms of its source, chemi- been covered by reviews of the results of various ap- cal nature and function in marine environments, from proaches, trying to understand the dynamics and chemi- early in the 20th century. cal nature of DOM (Trumbore and Druffel, 1995; Guo Since homeostatic feedback among the reservoirs of and Santschi, 1997; Nagata, 2000; Williams, 2000; bioelements has controlled the past global environments Kepkay, 2000; Myklestad, 2000; Ogawa, 2000; Benner, over the geological time scale (e.g., Berner, 1989), the 2002; Hansell and Carlson, 2002; Hedges, 2002). Some role and dynamics of dissolved organic carbon (DOC) fundamental new insights and observations have been have become of greater interest in the present global car- booked recently, which we have summarized in this re- bon cycle (e.g., Siegenthaler and Sarmiento, 1993). Rec- view. We thus focus here on papers published since 1995, ognition of the importance of the microbial loop has also although references to earlier fundamental work are in- has given us new insight into the role of DOM in marine cluded. ecosystems (e.g., Pomeroy, 1974; Azam et al., 1983). DOM has various functions and plays important roles Primary production is the ultimate source of organic mat- in chemical, biological and even physical oceanography. ter in the sea, but living biomass forms less than 1% of For example, DOM interacts with trace metals or total organic carbon in seawater, while more than 90% of radionuclides and controls their dynamics, it fuels the microbial loop, generates gases (CO, CO2) and nutrients with biological and photochemical reactions, absorbs and * Corresponding author. E-mail: [email protected] extinguishes light, and affects satellite images, etc. The Copyright © The Oceanographic Society of Japan. terrestrial input of DOM is also an important topic in the 129 global carbon budget as well as carbon dynamics in Thereafter, the HTC method was reassessed, im- coastal environments. In this review, however, we focus proved, and reborn as a highly precise method, suitable on recent advances in our understanding of the dynamics for measuring the seawater DOC (Sharp, 1997). In addi- and chemical characterization of oceanic DOM. Functions tion, the international distribution of certified reference of DOM in the environment are not described. materials (CRMs) for DOC analyses, including low car- bon water and deep seawater which were distributed by 2. What is DOM? Dr. Jonathan Sharp in 1997 and Dr. Dennis Hansell since The first difficulty that one must confront in any 1999, means that it has become possible to more correctly definition of DOM is to define the term “dissolved”. It compare DOC values measured by different workers. usually has an operational definition as any material that Consequently, both the accuracy and the precision of DOC passes through a given filter is termed “dissolved”. Glass data have been improved. Details of DOC measurement fiber filters (Whatman GF/F, with a nominal pore size of can be found in the recent review by Sharp (2002). 0.7 µm) are widely used for the collection and analysis A typical range of error of the DOC measurement by of organic substances. However, vastly different filter the recent HTC technique is 1–2% as a relative value, types have been used for study purposes. In reality, a spec- i.e., coefficient of variation (c.v.), or 0.5–1 µM as an ab- trum of material sizes exists; some material, such as col- solute value, i.e., standard deviation (s.d.) (Qian and loidal matter, does not fit neatly into either category. Mopper, 1996; Hansell et al., 1997b; Ogawa et al., 1999). Small-sized plankton pass through the filter; for exam- This is approximately 1/10 the error of the traditional ple, 22–38% of the total bacterial biomass (Lee et al., WCO method. The improved precision of the DOC 1995) and 100% of viruses, if they occur alone, are not method contributed greatly to the resolution of small dif- retained on the GF/F filter. We have to be aware that “dis- ferences in DOC concentrations in seawater in time and solved organic matter” does not exactly indicate the true space, although the previous method had been able to “dissolved phase” of organic matter in the sea. detect part of them qualitatively. This improvement was essential for the advancement of oceanic DOC study since 3. Distribution and Turnover Time of DOC in the we had to resolve the active but small pool of DOC rela- Ocean tive to the major part of DOC that was fairly refractory in seawater (see below). The introduction of this improved 3.1 Methodology of DOC measurement HTC method supplied quantitative and detailed informa- The abundance of DOM has generally been deter- tion for an understanding of the role of DOC in the oce- mined as dissolved organic carbon (DOC), which is a anic carbon cycle. major element of organic matter. A controversy about the measurement of DOC concentrations in seawater was 3.2 Distribution, accumulation and export of DOC in the current during the late 1980s and into the early 1990s. upper water column The high temperature (catalytic) combustion (HTC) Since the WCO method was established (Menzel and method introduced by Sugimura and Suzuki (1988) led Vaccaro, 1964), it has been consistently observed that to an argument about the existence of a much larger res- elevated concentrations of DOC appear in the upper wa- ervoir of DOC in the ocean than had been estimated by ter column, decreasing with increasing depth, followed the conventional method based on wet chemical oxida- by a low, uniform value in the deep water (e.g., Barber, tion (WCO, Menzel and Vaccaro, 1964). This problem 1968; Ogura, 1970). This finding suggests that DOC is excited many ocean scientists in the world because it had supplied by biological production in the surface ocean the potential to drastically change our concept of the oce- and consumed by microbial respiration, and the refrac- anic carbon cycle (Williams, 1992). After 1993, the con- tory component resistant to microbial degradation remains troversy calmed through further examination of the HTC in the deep water (Barber, 1968). In addition, recent ob- technique by a few communities (Hedges et al., 1993; servations using the high-precision HTC method were Sharp et al., 1993, 1995) and independent groups (Ogawa successful in detecting the detailed distribution and fluc- and Ogura, 1992; Tanoue, 1992; Cauwet, 1994). It was tuation of DOC within the surface water column. The found that the HTC measurement included a potentially observed changes in the surface DOC appear to be mainly high system blank relative to the DOC level in seawater caused by hydrological transport combined with biologi- (e.g., Williams, 1992; Benner and Strom, 1993). In con- cal production since they were observed over different clusion, the DOC values obtained from the HTC method productive sites along with vertical convection or hori- in the early stage were retracted (Suzuki, 1993) and the zontal advection of seawater (Copin-Montégut and Avril, concept of “new” DOC that might have represented a huge 1993; Carlson et al., 1994; Peltzer and Hayward, 1996; oceanic carbon reservoir, missed by the WCO method, Hansell and Carlson, 2001a, b). The apparent turnover of was rejected. DOC that accumulated and disappeared in the surface 130 H. Ogawa and E. Tanoue Fig. 1. Typical vertical distributions of DOC in the ocean according to a: conceptual classification of biological reactivity, b: size distribution and c: chemical composition. At each depth the concentration of bulk DOC is representative of the oceanic value reported after 1994, mainly including equatorial, subtropical and temperate zones (see Table 1), without the Antarctic and the Arctic oceans.
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