Preferred Citation: . The Oceans, Their Physics, Chemistry, and General Biology. New York:  Prentice-Hall,  c1942 1942.

Organisms and the Composition of Sea Water

Chemical Composition of Marine Organisms

Alterations of the concentration of the dissolved constituents of sea water are brought about by the development and subsequent death and disintegration of organisms. Virtually all of the substances extracted from the water are returned to solution by metabolic processes or by disintegration of the organisms, but the elements removed are returned to solution at some later time and often in some other part of the water column. Hence, the modifications may be in opposite directions at different times and localities. A small fraction of the organic remains accumulates on the sea bottom and is lost to the cycle.

Sea water probably contains in solution all of the chemical elements, although only some fifty have yet been detected. There is a large amount of data on the occurrence of various elements in marine plants and animals, but unfortunately the material is far from complete for any one biological group. Either a few elements only have been determined—such as iodine, for example, which has been thoroughly investigated—or only a portion of the organisms—for example, the skeletal structures—has been analyzed.

Vinogradov (1935, 1937) has compiled the chemical analyses of the lower plants and animals, both aquatic and terrestrial. He reports some sixty of the elements that have been found in one or more species. Webb and Fearon (1937) have tabulated thirty-nine elements that are commonly found and have divided these into two groups according to their apparent importance to living things: (1) eighteen invariable elements, and (2) twenty-one variable elements. These classes are further sub-divided on the basis of the concentration in which the elements are present. Seven elements are listed as contaminants (table 46).

The primary invariable elements are the essential constituents of carbohydrates, lipides (fats), and proteins. Some of the invariable elements classed as secondary or as microconstituents are always present in the lipides and proteins. This list is for plants and animals in general and not for marine forms alone. Comparison of tables 46 and 36 shows that nine elements (starred) detected in organisms have not yet been

reported for sea water, and that seven (uranium, thorium, cerium, lanthanum, yttrium, scandium, and radium), known to occur in sea water, are not listed by Webb and Fearon. Radium, at least, should be added to their list (p. 184).

Invariable (18) Variable (21) Contaminants
Primary 1–60% Secondary 0.05–1% Microconstituents <0.05% Secondary Microconstituents

* Not yet reported for sea water.

Hydrogen Sodium Boron Titanium[*] Lithium Helium
Carbon Magnesium Fluorine Vanadium Beryllium[*] Argon
Nitrogen Sulphur Silicon Zinc Aluminum Selenium
Oxygen Chlorine Manganese Bromine Chromium[*] Gold
Phosphorus Potassium Copper Cobalt[*] Mercury
Calcium Iodine Nickel Bismuth[*]
Iron Germanium[*] Thallium[*]

The lack of comparable data for the different types of organisms makes it necessary to consider their composition under three headings—namely, organic material (largely carbohydrates, lipides, and proteins), inorganic skeletal structures, and inorganic solutes in the body fluids. Although the proportions of carbohydrates, lipides, and proteins may vary considerably, the composition of any one type is rather constant, so that the average values in table 47 can be used with some confidence. Furthermore, there are numerous determinations of lipides (ether extract) and protein (based on nitrogen determinations), and from these measurements and the loss on ignition the carbohydrate may be computed. Skeletal structures differ so much in composition and in their mass, compared to that of the organic material, that they must be considered separately. Inorganic solutes in the body fluids are considered as a separate class, because they apparently do not differ very much in composition

and concentration from the surrounding sea water; consequently, the presence of water in the original sample will give merely rather high values for the inorganic solutes. The great changes in relative composition are found in the organic material and skeletal structures, for which reason, if a mixed plankton sample containing sea water is analysed, the results for the organic material and skeletal structures will not be materially affected if it is considered that all the sodium or chloride, as well as other elements in the proportions in which they occur in sea water, are in the body fluid or sea water.

Percentage composition Relative proportions by weight, C = 100
Element Carbohydrates Lipides Proteins Element Sea water Lipides Proteins
O 49.38 17.90 22.4 C 100 100 100
C 44.14 69.05 51.3 P 0.05 3.1 1.4
H 6.18 10.00 6.9 N 0.5 0.88 34.7
P 2.13 0.7 S 3150 0.45 1.6
N 0.61 17.8 Fe 0.07 0.2
S 0.31 0.8
Fe 0.1

In table 47 are given the average compositions of the three great classes of organic material (Rogers, 1938) and the relative proportions in which their component elements occur in sea water. The oxygen and hydrogen are not considered, and the values are adjusted to C = 100. The values for C, S, and Fe are from table 36; those for N and P are the average winter values in the English Channel (pp. 252, 258). In the lipides, phosphorus is concentrated, and in the proteins the nitrogen and phosphorus show a great increase with respect to carbon. The fact that sulphur, which is one of the relatively abundant elements in sea water, is a minor constituent of the lipides and proteins in organic material indicates that carbon, here used as the reference element, is itself markedly concentrated. The values given in table 47 are general averages, and those for marine organisms may differ slightly. It should be noted that changes in the proportions of carbohydrates, lipides, and proteins will modify the ratios in which the above-mentioned elements will be removed from the water. Many of the other elements that are concentrated by organisms—for example, iodine, iron, and copper—probably form a part of the organic material or they occur in the skeletal structures, as it is difficult to see how the free ions could be retained in the body fluids

at such tremendously higher concentrations than those in the surrounding sea water.

In table 48 are given analyses of certain types of skeletal material. In each case there is some organic matter, which is highest in the lobster carapace, and even in the phosphatic brachiopod shell it forms a large fraction. Of course, there are wide ranges in the proportion of inorganic skeletal structures in the whole organism, and in some cases such structures may be entirely lacking. The first three examples are for calcareous types with CaCO3 predominating, but in some groups MgCO3 forms an important part of the shell. The lobster may be considered as representative of the arthropods in general, although the proportion of organic matter is probably even greater in the small forms. The amount of phosphate is notable in the lobster and even more so in the phosphatic brachiopod shell, which is predominantly calcium phosphate. The sponge spicules are virtually pure hydrated silica and may be taken as representative of the diatom and radiolarian skeletons. The silica, iron, and aluminum in the other analyses probably represent impurities introduced by the presence of clay and sand grains. These analyses cannot be regarded as complete, and further examination will undoubtedly reveal many other elements present in small amounts. It should be noted that chlorine and sodium, the two most abundant elements in sea water, are not shown in any of these analyses. These elements form soluble compounds and hence would not be suitable for skeletal structures. From table 48 it can be seen that the development or re-solution of skeletal structures of marine organisms may be expected to affect the concentrations

of calcium, magnesium, carbon (as carbonate), sulphur, phosphorus, and silicon. Because of their relatively greater abundance in sea water, the distribution of magnesium and sulphur cannot be expected to show any appreciable effects of biological activity, but the distributions of the other elements mentioned do show certain features that can be attributed, in part at least, to the fact that they are important constituents of skeletal structures.

PERCENTAGE COMPOSITION OF SKELETAL MATERIAL (Recalculated from Clarke and Wheeler, 1922)
Substance Foraminifera (Orbitolites marginatis) Coral (Oculina diffusa) Calcareous alga (Lithophyllum antillarum) Lobster (Homarus sp.) Phosphatic brachiopod (Discinisca lamellosa) Siliceous sponge (Euplectella speciosa)
Ca 34.90 38.50 31.00 16.80 26.18 0.16
Mg 2.97 0.11 4.36 1.08 1.45 0.00
CO3 59.70 58.00 62.50 22.40 7.31 0.24
SO4 0.68 0.52 4.43 0.00
PO4 tr tr tr 5.45 34.55 0.00
SiO2 0.03 0.07 0.04 0.30 0.64 88.56
(Al,Fe)2O3 0.13 0.05 0.10 0.44 0.32
Organic matter, etc 2.27 3.27 1.32 53.45 25.00 10.72

The relative concentrations of the elements that are abundant in the body fluids do not differ very much from those in sea water (table 49). Although little is known concerning the less abundant elements, it appears that the inorganic portion of the body fluids can be considered as slightly altered sea water. Therefore, this part of the organism cannot play any appreciable part in modifying the composition of the water. Although the composition and concentration of the inorganic solutes is of no particular importance in the present problem, these features have been studied intensively in problems of osmotic pressure relations (chapter VIII) and in connection with the mechanism of solute and water exchange between aquatic organisms and their environment. These fields have been reviewed by Rogers (1938).

RELATIVE COMPOSITION OF BODY FLUIDS (Adjusted to Na = 100. Data from Robertson, 1939)
Element Sea water Echinus esculentus (Sea urchin) Homarus vulgaris (Lobster) Cancer pagurus (Crab)
Cl 180 182 156 156
Na 100 100 100 100
Mg 12.1 12.0 1.5 5.7
S in SO4 8.4 8.5 2.2 6.7
Ca 3.8 3.9 5.0 4.8
K 3.6 3.7 4.7 4.0

Thus far only the various fractions of the organisms have been discussed, and it is of interest to consider the composition of the entire plant or animal. As the plants are the primary “consumers” of inorganic material, it would be desirable to know the composition of such important groups as the diatoms and peridinians, but no complete analyses of these forms have been made. What information we have will be discussed below. The data for animals is also far from complete, but in table 50 are given three examples. The relative compositions have been adjusted to Na = 100, and, for comparison, the constituents of sea water are given in the same way. The relatively high proportions of the elements abundant in sea water which occur in the copepod analysis indicate the presence of considerable sea water in the original sample. As Archidoris possesses internal calcareous structures, the calcium content is high. Consequently the proportions of the elements constituting the organic material are rather low in these two cases. It is immediately obvious, however, that the essential constituents of the organic material, such as carbon, nitrogen, and phosphorus, are very high when compared to their relative concentrations in sea water.

Element Calanus (Copepod) Vinogradov, 1938 Fish[a] (Average) Archidoris britannica (Nudibranch) McCance & Masters, 1937–38 Sea water Concentration factors
Copepod Fish Nudibranch

aFrom analysis of Jowett and Davies (1938), Clements and Hutchinson (1939).

bWinter values, English Channel.

cHigher values from table 36.

Cl 194 180 180 1.1 1.0
Na 100 100 100 100 1.0 1.0 1.0
Mg 5.6 36 156 12.1 0.46 3.0 12.9
S 25.9 259 7.1 8.4 3.1 31 0.85
Ca 7.4 52 262 3.8 1.9 13.7 69
K 53.7 383 20 3.6 15 109 5.5
Br 1.7 0.6 3
C 1113 ca 4100 ca 480 0.26 4,300 15,800 1,850
Sr 11 0.12 92
Si 1.3 0.001[b] 13,000
F 69 0.01 6,900
N 280 1276 107 0.001[b] 280,000 1,276,000 107,000
P 24.1 256 6 0.0001[b] 241,000 2,560,000 60,000
I 0.04 0.0005 80
Fe 1.3 1.3 0.23 0.0002[c] 6,000 6,000 1,000
Mn 0.0008 0.0001[c] 8
Cu 0.008 0.43 0.0001[c] 80 4,300


If the relative amounts of the various elements in these animals are divided by their relative concentrations in sea water, a series of concentration factors, referred to sodium, are obtained. It will be seen that chlorine would give virtually identical results. The concentration factors range from about unity up to over two million for phosphorus in the fish, and in all three cases they are greatest for nitrogen and phosphorus. If it is assumed that the rates of diffusion of all substances and their rates of absorption by the organisms depend only upon the amounts of the ions in the water, then the concentration factors should be a measure of the time required to accumulate them. Those elements having the highest concentration factors would then be the ones that might limit the rate of growth. The data in table 50 indicate that nitrogen and phosphorus may very well be limiting elements in the sea, although it should be remembered that the examples are for animals that must obtain their supply of these elements either directly or indirectly from the plants. If the total carbon in sea water had been used as the reference element, only nitrogen and phosphorus would have significantly larger factors. But it is obvious that carbon is itself concentrated more than one thousandfold with reference to the major elements in sea water. According to table 50 the relative concentration

in sea water of the following elements may be markedly affected by biological activity: carbon, silicon, fluorine, nitrogen, phosphorus, iron, and copper. Other elements might be included if the analyses were more complete or if other types of organisms were examined.

Element Sea water Diatoms Peridinians Copepods Concentration factors (referred to carbon)
Diatoms Peridinians Copepods

aWinter values, English Channel.

bHigher value, table 36.

C 100 100 100 100 1 1 1
N 0.5[a] 18.2 13.8 25.0 36 28 50
P 0.05[a] 2.7 1.7 2.2 54 34 44
Fe 0.07[b] 9.6 3.4 0.13 137 49 2
Ca 1420 12.5 2.7 0.66 0.01 0.002 0.0005
Si 0.4[a] 93.0 6.6 0.13 232 16 0.3

In table 51 are given the relative concentrations of certain elements in diatoms, peridinians, and copepods, adjusted to C = 100. The data for the photosynthetic forms are recomputed from Vinogradov (1935), and for copepods are the same as those given in table 50. The concentration factors for nitrogen and phosphorus are about the same in all three forms. In the diatoms, iron is higher, while silicon has the highest factor, which may indicate that these elements also limit the rate of growth. For the peridinians the factors for nitrogen, phosphorus, and iron are nearly the same.

Organisms and the Composition of Sea Water

Preferred Citation: . The Oceans, Their Physics, Chemistry, and General Biology. New York:  Prentice-Hall,  c1942 1942.