THE ENVIRONMENTAL CONDITIONS OF PLANTS IN ARID REGIONS.
The environmental conditions encountered by plants in the arid regions are widely different from those of moister regions. Precipitation is not only slight, but it shows an enormous range in variation from year to year; the rate of evaporation is high; the temperature of the air and soil varies widely both during the day and with the seasons; the light is of great intensity, and the soil is low in humus content and may contain an excess of salts. These, the most striking physical factors of deserts, are present in different combinations with resultant differences among deserts, and an arid region may be so large as to include such variation within its borders. Also, a desert may be so far from the ocean, or it may include such diversity of topography, as to show great variation in biological features as well as in its surface phenomena.
In the flora of any arid region, the mutual relations of the constituents of the flora and its general and detailed relation to the physical environment are quite different in the main from these features in the flora of the more humid regions. Thus, the leading relation is the relation to water, and on the response of the plants to this relation much of the phenomena associated with plant life in the desert directly depends. For example, in the extreme deserts it is probable that the elements of competition between the perennials, which is an important factor in the survival of a species in the moister regions, is wholly lacking. It should be noted, however, that in the less extreme deserts, as in the vicinity of the Desert Laboratory, competition exists between plants, although this is not at first apparent. In this case the competition is not for room, but for water, and is not manifest by palpable crowding, but by invisible competition of the roots. Thus the reactions are with the physical environment and are exhibited in a variety of ways, some of which concern the plants themselves in an intimate manner, being morphological and physiological, some being concerned with the flora as a whole. The environmental responses are often obscure and complex, but in other instances they are less obscure and apparently direct.
As is well known, a desert flora is in part perennial, lasting with but little outward change from season to season, and in part ephemeral, consisting of short-lived species which appear with the rains and which disappear with the return of the dry season. The ephemeral flora differs in no essential regard from annuals of the moister regions; also, the environment to which they are exposed closely resembles the environment of the annuals of such regions. But the perennial desert flora, on the other hand, offers very striking departures from the corresponding flora of the moister regions, just as the environment to which they are exposed for the most of the year is also different. It will be sufficient, for the purpose of bringing out the point of view, to notice a few of the leading characteristics of desert plants and of their physical environment and, in a few instances, to observe possible relations between the two.
The most obvious features of desert plants are associated, in whatever way, with the subaerial portions. Leaves are usually greatly reduced or wanting, during the dry seasons at least. Spines are frequently present and the exposed parts are often well covered with hairs. The stomata are sometimes deeply sunken, the cuticle often very heavy, and a waxy substance may cover leaves or stems. The chlorophyll-bearing cells are arranged with the long axis at right angles to the leaf or stem surface. All or most of these characters are associated with the low humidity of the air. In certain deserts plants are also to be found with greatly enlarged stems and branches which serve as water-storage organs. It should be noted, however, that plants which do not have a constant surface undergo many marked changes with a betterment of the water relations, particularly if this comes when the temperatures are favorable. For example, many cacti organize leaves which are unsuited in structure for periods of extreme drought, and which consequently fall away soon after the close of the rainy season. These leaves enormously increase the rate of transpiration at a time when this is not injurious.
It is not in the subaerial parts alone, however, that the plants of one desert are different from those of another, that plants are unlike in the same desert, or that plants of a desert are different from those of the more humid regions. The root-habits also exhibit not a little diversity and show marked reactions to the pressure of their environment. For instance, the desert shrubs of the region surrounding the Desert Laboratory at Tucson have well-marked root-systems, apparently constant under natural conditions, which may roughly be designated as the tap-root type, the superficial type, and the generalized type. Other conditions being equal, species with characteristic root-types have also characteristic distribution, or exhibit in other regards consistent reactions. Thus, the widely extending and superficial type of roots is confined, among independent plants, to such as have water-storage capacity. The relation of this root-type to the distribution will be mentioned later. Plants with a dominating tap-root are confined to areas where the soil is relatively or actually deep, while species having generalized roots have a local distribution which may be considered the maximum.
The relation of the superficial type of root-system to the distribution of the species is not so apparent in this relation in plants as with the other types of roots, but is undoubtedly close and possibly definitive. The absorption roots of plants with water-balance mostly lie less than 10 cm. beneath the surface and are thus subject to extreme desiccation for the maximum time, or in other words they are exposed to favorable moisture conditions the minimum time. How long the optimum water-absorption time may be for such species is not known, but that it is longer than might be supposed (from the period certain species can survive without water) is highly probable from the facts concerning their distribution. In brief, the best development of the fleshy species in question occurs where the rainfall is periodic, occurring twice each year, and they are wanting or sparse where the rainfall is uncertain or occurs but once annually. Had these plants a deeply penetrating type of root-system the local as well as the general distribution would be very different from what it now is. Owing to the unfavorable character of the rainfall in southern Algeria, plants with a water-balance are wanting there, just as they are wanting in portions of southwestern United States where the amount or the character of the precipitation is likewise unfavorable.
Besides the characteristics of the root-systems of the desert perennials whose significance has been sketched, there are other features of importance. For example, the roots lying near the surface bear tufts of delicate roots, which are formed during periods of active growth and perish when such seasons cease. By the organization of deciduous rootlets the absorption area of the species is enormously increased, and quickly, without at the same time increasing the distance of water transport.
There need be mentioned here only one additional feature of the desert plants. It is now known, in brief, that the non-fleshy perennials of the desert, not halophytes, may possess a very dense cell-sap. This fact has been demonstrated in the subaerial parts of several species, and is assumed to hold for the roots also of the same species. As suggested, not all of the desert plants, however, are capable of developing dense juices. Thus, certain fleshy species, and such mesophytes of the desert as have been studied in this connection, do not have more highly concentrated cell-sap than the ordinary plants of the humid regions. Further, it appears that desert species which, under natural conditions and during the dry seasons, form extremely dense juices, lose this capacity when grown under humid conditions. So far as is now known, species capable of developing a cell-sap with high osmotic power have generalized roots, although this may be of no especial significance; but the relation of this capacity to survival in an arid substratum is apparent and vital.