Disease in Plants. Ward Harry Marshall

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СКАЧАТЬ in a 15 per cent. solution of calcium chloride developed no root-hairs, though they formed in a 5 per cent. solution, and were very numerous in a 0.5 per cent. solution, or in water alone. In a 10 per cent. nutritive solution the plants developed no root-hairs, though they were abundant in a 1 per cent. solution.

      Wheat plants with their roots in a 15 per cent. solution of potassium nitrate bore no root-hairs, but they were numerous in a 2 per cent. solution of the same salt.

      These are extreme cases, for, although the roots were not killed, they were strongly inhibited in their growth by the more concentrated solutions. However, experiments of this kind at least bring vividly before us what variations are possible, and suggest that similar events on a smaller scale may occur in a soil which yields large quantities of soluble substances, e.g. when freshly manured. Obviously these facts have a practical significance as regards kind of soil, drainage, season (e.g. drought or wet), etc.

      But there are other factors which rule the development of root-hairs, and some experiments by Lesage show that the correlations between the development of root-hairs and roots are probably much more complex than had been suspected; for he finds that if the lateral rootlets of a Bean, in a water culture, are suppressed, the main rootlet develops numerous and very long hairs to compensate the loss in surface, a matter of obvious importance in the discussion of cases where roots have been injured in the soil.

      Before proceeding further it is necessary to look a little more closely into the structure of a single hair.

      It is a tubular prolongation of a single cell of the external covering of the young root, usually about 1 to 3 mm. in length, and 0.01 to 0.10 mm. in diameter. In special cases the root-hairs of some water plants may reach 5 to 18 mm. in length, but of course I am referring to the ordinary land plants of agriculture and forestry. This tubular prolongation is closed and rounded off at the distal free end, and opens at the proximal end into the cell of which it is a protrusion.

      The whole structure is bounded by an extremely delicate and elastic wall of cellulose, which Frank says is of special composition, almost too thin to measure in many cases, but often somewhere near 0.005 to 0.001 mm. in thickness. This thin membrane is remarkably permeable by water, or dilute solutions, as is shown by the rapidity with which a root-hair collapses if exposed to evaporation, or with which dense solutions abstract water from it, or with which solutions may be seen to penetrate it under the microscope.

      Overlying the thin cell-wall proper, on the outside, is a thin gelatinous layer, a product of alteration of the outermost lamellæ of the former.

      Closely lining the proper cell-wall on the inside, is an extremely thin layer of living protoplasm, and somewhere in this protoplasm is a distinct cell-nucleus.

      The interior of the tube is filled with cell-sap, and it is the osmotic pressure of this cell-sap which keeps the whole living instrument tense and rigid, and the thin protoplasmic film close pressed against the cellulose cell-wall.

      Nothing whatever can pass into the cell-sap, or out from it, without traversing both the lining of living protoplasm and the cell-wall.

      If we gently pull a living root, of wheat, pea, mustard, etc., from a normal soil, we find particles of soil so closely adherent to the root-hairs that they cannot all be washed off without tearing the hairs: the root-hairs establish relations of contact with these particles, so close that they are cemented to the solid surfaces by means of the gelatinous layer already referred to. This peculiarity has the following consequences. In the first place, the enormous holdfast, ensured by the millions of points of adherence, enables the plant to withstand even powerful lever actions from above, and provides fixed points against which the root-tips can work as they drive deeper into the soil. In the second place, the intimate contact of the root-hairs and particles of soil, ensures that the films of water held by surface-action on the soil-particles and root-hairs shall be in continuity with the water saturating the cell-walls of the latter, and therefore with the protoplasm and cell-sap in their interior. The importance of this at periods when the soil is "dry" will be obvious, when we reflect that no soil is ever naturally so dry that surface-films of water are absent from the particles.

      The fact that the root-hair contains living protoplasm, enables us to understand to a certain extent the results of the following experiments.

      If we have a leafy and healthy plant, with roots, bearing numerous root-hairs, properly established in suitably moist soil in the pot, the roots cease to absorb water if the temperature of the soil falls below a certain minimum, though they recommence to do so if the temperature is raised again: this has nothing to do with the temperature of the upper parts of the plant, or of the air, and the latter may be so high that the plant rapidly droops from loss of water at the leaves, which is not being compensated owing to the inactivity of the roots.

      Similarly we may have the air so cold, at a time when the soil is warm enough to keep the root-hairs actively at work, that the plant becomes surcharged with water, which escapes from the leaves like drops of dew. The temperatures necessary to cause these disturbances in the action of the living root-hairs vary for different plants, and even for different varieties of the same species.

      Similar arrestation of the functions of the roots may be brought about by removing the oxygen from the soil around the root-hairs, and replacing it by carbon-dioxide, or the vapour of chloroform. If not kept too long in such a condition, the plant recovers rapidly on admitting atmospheric oxygen, which is always present in a normal well-drained soil both as gas in the capillary interspaces, and dissolved in the water on the surfaces of the particles. If the access of oxygen is delayed, however, as often happens in rainy seasons and in wet soils, the root-hairs are killed, and rot sets in. A good instance of this has lately been given by Heinricher in the case of potatoes.

      Notes on Chapter V

      For the further pursuit of this subject the reader should consult Sachs' Lectures, II. and XV.; Sorauer, A Popular Treatise on the Physiology of Plants, 1895, chapters II. and IV., and Pfeffer's Physiology, pp. 149-163. The principal paper on root-hairs referred to in the text is Schwarz, "Die Wurzelhaare der Pflanzen," in Unters. aus dem bot. Inst. zu Würzburg, I. Heft 2, 1883, p. 140, where a very exhaustive account of these organs will be found.

      CHAPTER VI.

      THE FUNCTIONS OF ROOT-HAIRS

      Excretions from root-hairs—Osmotic phenomena—Turgescence—Plasmolysis—Control of the protoplasm in absorption, etc. Selective absorption.

      We see then that the root-hairs are the active living instruments in absorbing the water (containing small quantities of dissolved substances) of the soil.

      If the living root-hairs are so numerous and so active, however, a natural inference is that they must exert some influence on the composition or arrangement of their environment. All the teachings of modern physiology go to show that such a living cell as I have sketched cannot carry on its life, brief though it be—the root-hairs are active for about four or five days—without forming substances of the nature of excreta, and we should expect some of these to pass out to the soil.

      Sachs showed, in 1860, that roots growing in contact with polished marble corrode the surface of the mineral, and Nobbe, in 1876, showed that the roots of seedlings reduce potassium permanganate, a fact which Molisch confirmed in 1887. The latter observer also proved that living root-hairs secrete substances which colour a solution of guaiacum blue, oxidise pyrogallic acid and other organic substances, and rendered it probable that they excrete some substance which inverts cane-sugar, and in some cases even small quantities of a diastatic enzyme.

      Molisch also confirmed an old observation, that roots excrete carbon-dioxide; and he and Czapek showed СКАЧАТЬ