Temperature

Because of the close association of temperature with other factors, we have already had numerous occasions to refer to it in the course of discussion. It is certainly one of the most important of physical factors. However, it is not easy to generalize and designate any one factor as being more important than any other in a complex natural environment where all of the numerous known factors operate simultaneously. According to Liebig's law of the minimum, production is limited by the factor occurring in minimal quantity. This factor is obviously not the same for each circumstance. Hence, actually, the factor occurring in minimal quantity, whatever it may be, is temporarily the most important factor. It becomes the so-called “limiting factor” which has been defined as “any of the direct factors which is not optimal at the conditions present in a water mass, viz., without any change of the other factors.”

The rate of metabolism, hence the rate of growth and reproduction of any given species, is regulated by temperature. Just what will be the required range of temperature is dependent upon the species, but the warmest as well as the coldest waters of the sea are able, when other external factors are favorable, to produce a characteristic type of flora

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adapted to the conditions. The separate species are either eurythermic or stenothermic, depending upon whether their tolerance to temperature range is great or small. Different genera and different species within a genus may indicate a temperature preference, as shown by the following maximal growth periods for certain species in the Gulf of Maine at the surface temperatures indicated (Gran and Braarud, 1935):

Neritic species of the higher latitudes are also durothermic with respect to low temperatures during resting stages. These resting stages are very resistant to extreme temperatures and have been found to be viable even though they have spent unfavorable periods frozen solidly in ice.

We may conclude, then, that one great influence of temperature is in determining the character, that is, the composition, of species that will develop in any one region at various periods of the year. In temperate latitudes it is possible, depending on season and other conditions, for both high-latitude and subtropical species to grow provided either that “seed” can be carried in by currents at the proper season or that durothermic resting stages can survive the “off” season and germinate again with return of the growing season.

In this way Cleve's geographical diatom types, namely, arctic, boreal, temperate, and tropical, are explained not as having been produced in and carried from these separate areas but rather as having their metabolic requirements adjusted to temperatures characterizing these regions. During seasons of unfavorable temperatures the stock of separate types is maintained as resting-spores or resistant individuals with potentialities of quick response upon the return of their specific temperature tolerance, the different species succeeding each other in order according to their requirements.

Perhaps one of the most far-reaching influences of temperature is indirect through its effect on the viscosity of the water. We have had frequent need to mention how vitally important it is that the open-sea plants be kept within the euphotic layer. The degree of viscosity or internal friction of the water is vital to passively floating organisms with a specific gravity greater than the water in which they live. Variations

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in salinity have only a small influence on viscosity, and owing to the small variations in salinity, except near shore, viscosity is not materially affected from this cause; but a change in temperature from 25°C to 0°C is sufficient to double the viscosity. With this in mind, we see how important it is to phytoplankton suspension when a transition or thermocline with a marked temperature gradient occurs within the euphotic zone. The rate of sinking at the transition layer may be sufficiently retarded to allow an accumulation of plants that would otherwise not have occurred owing to reduction of stock by sinking, with the subsequent slowing up of production that accompanies such withdrawal of stock (p. 772).

Winter and summer variations which occur in the structure of certain species are attributed to the temperature variations of the seasons in areas where there is great temperature range, but temperature differences in higher and lower latitudes have the same effect, consequently summer forms of high-latitude species may occur in the warmer seas or vice versa. The structural differences are concerned mainly with adaptations leading to a better adjustment to the floating existence. The summer or warm-water forms have thinner shells and are of more slender build. A classical example is given in the diatom Rhizosolenia semispina (fig. 70), wherein a temperate form (semispina) and an arctic form (hebetata), once regarded as separate species, may originate one from the other or wherein opposite ends of a single cell may even show characteristics of both. The form hebetata is now regarded as a primitive resting stage of the form semispina.

In concluding this discussion it should be emphasized that even though the conditions with respect to light, temperature, nutrients, and so forth may be favorable to diatom production, yet no spontaneous population development can take place if previous conditions with respect to any factor have left the water sterile of vegetative plants or of resistant resting spores of species capable of taking advantage of the return of good conditions. This may explain in some instances the dearth of phytoplankton in offshore situations where the great depth of water prohibits the ready return of resting stages common to species of neritic waters. The phenomenal outbursts of diatoms in arctic regions coincident with the spring melting of ice have been explained as probably being associated with the rapid germination of spores that have been locked in the ice and are released through melting. It is known that these regions simulate biological conditions of coastal areas and the population is largely neritic. In investigating oceanographic conditions at a line of stations through Barents Sea from the coast northward to the Arctic ice, Kreps and Verjbinskaya (1930) found in spring that the early vernal flares of diatoms were confined to coastal waters and to waters near the Arctic ice, while between these and at some distance from

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shore in branches of the North Cape Current of Atlantic water, the population was poor although nutrients were high. In the Arctic waters near the front of melting ice both nutrients and diatoms were abundant.

The stimulation of phytoplankton growth commonly observed to be coincident with the mixing of two bodies of water with different characteristics may also result from an inoculation of a sparse population living under poor conditions into more favorable waters which have lacked spores suitable to take advantage of the good physical-chemical environment.

Population control through biological factors, such as animal consumption, is discussed in chapter XVII.