Changes occurring during Germination.—When a seed is placed in the soil under favourable circumstances, it becomes the seat of an important and remarkable series of chemical changes, which result in the production of the young plant. Experiment and observation have shown that heat, moisture, and air, are necessary to the production of these changes, and though probably not absolutely essential, the absence of light is favourable in the early stages. The temperature required for germination varies greatly in different seeds, some germinating readily at a few degrees above the freezing point, and others requiring a tolerably high temperature. The rapidity with which it takes place appears to increase with the temperature; but this is true only within very narrow limits, for beyond a certain point heat is injurious, and when it exceeds 120° or 130° Fahrenheit, entirely prevents the process. The presence of oxygen is also essential, for it has been shown that if seeds are placed in a soil exposed to an atmosphere deprived of that element, or if they be buried so deep that the air does not reach them, they may lie without change for an unlimited period; but so soon as they are exposed to the air, germination immediately commences. Illustrations of this fact are frequently observed where earth from a considerable depth has been thrown up to the surface, when it often becomes covered with plants not usually seen in the neighbourhood, which have sprung from buried seeds. When all the necessary conditions for germination are fulfilled, the seed absorbs moisture, swells up, and sends out a shoot which rises to the surface, and a radicle which descends—the one destined to develop the leaves, the other the roots, by which the plant is afterwards to derive its nutriment from the air and the soil. But until these organs are properly developed, the plant is dependent on the matters contained in the seed itself. These substances are mostly insoluble, but are brought into solution by the atmospheric oxygen acting upon the gluten, and converting it into a soluble substance called diastase, which in its turn reacts upon the starch, converting it first into dextrine, and then into cellulose, and the latter is finally deposited in the form of organised cells, and produces the first little shoot of the plant. At the first moment of germination, the oxygen absorbed appears simply to oxidize the constituents of the seed, but this condition exists only for a very limited period, and is soon followed by the evolution of carbonic acid, water being at the same time formed from the organic constituents of the seed, which gradually diminishes in weight. The amount of this diminution is different with different plants, but always considerable. Boussingault found that the loss of dry substance in the pea amounted in 26 days to 52 per cent, and in wheat to 57 per cent in 51 days. Against this, of course, is to be put the weight of the young plant produced; but this is never sufficient to counterbalance the diminished weight of the seed, for Saussure found that a horse bean and the plant produced from it weighed, after 16 days, less by 29 per cent than the seed before germination. The same phenomenon is observed in the process of malting, which is in fact the artificial germination of barley, the malt produced always weighing considerably less than the grain from which it was obtained. It was believed by Saussure, and the older investigators, that the carbonic acid evolved was entirely produced from starch and sugar; and as these substances may be viewed as compounds of carbon and water, the change was very simply explained by supposing that the carbon was oxidised and converted into carbonic acid and its water eliminated. But this hypothesis is incapable of explaining all the phenomena observed; for woody fibre, which is one of the chief constituents of the young plant, contains more carbon than the starch and sugar from which it must have been produced, and we are, therefore, forced to admit that the action must be more complicated. There is every reason to believe that the nitrogenous constituents of the seed are most abundantly oxidized, for they are remarkably prone to change; but the action of the air is not confined to them, and it appears most probable that all the substances take part in the decomposition, and the process of germination may, in some respects, be compared to decay or putrefaction, which, like it, is attended by the absorption of oxygen and evolution of carbonic acid; but while in the latter case the residual substances remain in a useless state, in the former they at once become part of a new organism.

Changes occurring during the After-growth of the Plant.—When the plant has developed its roots and leaves, and exhausted the store of materials laid up for it in the seed, it begins to derive its subsistence from the surrounding air, and to absorb carbonic acid, water, ammonia, and nitric acid, and to decompose and convert them into the different constituents of its tissues. These changes take place slowly at first, and more rapidly as the organs fitted for the elaboration of its food are developed. The roots and the leaves are equally active in performing this duty, the former absorbing the mineral matters along with the carbonic acid, ammonia, nitric acid, and moisture in the soil, or the manure added to it; the latter gathering the gaseous substances existing in the air. Each of these undergoes a series of changes claiming our consideration.

Decomposition of Carbonic Acid.—Carbonic acid, which appears to be absorbed with equal readiness by the roots, leaves, and stems, undergoes immediate decomposition, its carbon being retained, and its oxygen, in whole or in part, evolved into the air. This decomposition occurs only under the action of the sun's rays, and has been found to be proportionate to the amount of light to which the plant is exposed. It takes place only in the green parts of plants, for though the roots absorb carbonic acid, they cannot decompose it, or evolve oxygen; and the coloured parts, the flowers, fruits, etc., have an entirely opposite effect, absorbing oxygen and giving off carbonic acid. The absorption of carbonic acid and escape of oxygen has been proved by numerous direct experiments by Saussure and others, in which both atmospheric air and artificial mixtures containing an increased quantity of carbonic acid have been employed. Saussure allowed seven plants of periwinkle (Vinca minor) to vegetate in an atmosphere containing 7·5 per cent of carbonic acid for six days, during each of which the apparatus was exposed for six hours to the sun's rays. The air was analysed both before and after the experiment, and the results obtained were—

Volume of the air.Nitrogen.Oxygen.Carbonic Acid.
Before the experiment,574641991116431
After "5746433814080
————————
Difference,0+139+292-431

In this experiment the whole of the carbonic acid, amounting to 431 volumes, was absorbed, but only 292 volumes of oxygen were given off. Had the carbonic acid been entirely decomposed, and all its oxygen eliminated, its volume would have been equal to that of the acid, or 431, so that in this instance 139 volumes of the oxygen of the carbonic acid have been retained to form part of the tissues of the plant. On the other hand, the nitrogen is found to be increased after the experiment. It might be supposed that the nitrogen evolved had been derived from the decomposition of the nitrogenous constituents of the plant, but this cannot be the true explanation, because in this particular case it greatly exceeded the whole nitrogen contained in the plants experimented on. Its source is not well understood, but Boussingault supposes it to have existed in the interstices of the plant, and to have escaped during the course of the experiment. Saussure found that the oak, the horse-chesnut, and other plants, absorb oxygen and give off carbonic acid in less volumes than the oxygen, while the house-leek and the cactus absorb oxygen without evolving carbonic acid. The absorption and decomposition of carbonic acid takes place only during the day, and matters are entirely reversed during the night, when oxygen is absorbed and carbonic acid eliminated from all parts of the plants.

Although the action occurring during the night is the reverse of that which takes place during the day, it is in no degree to be attributed to a re-oxidation of the carbon which had been deposited in the tissues of the plant. It appears, on the contrary, to be a purely mechanical, and not a chemical process. During the night the sap continues to circulate through the vessels of the plant, and moisture, carrying with it carbonic acid in solution, is absorbed by the roots; but when it reaches the leaves, where the sun's light would have caused its decomposition during the day, it is again exhaled unchanged. The oxygen absorbed during the night must, however, take part in some chemical processes, for if it were merely mechanical, the absorption would not be confined to that gas alone, but would be participated in by the other constituents of the air. Moreover, the amount of absorption varies greatly in different plants—being scarcely appreciable in some, and very abundant in others. Plants containing volatile oils, which are readily converted into resins by the action of oxygen, or those containing tannin or other readily oxidizable substances, take up the largest quantity. This is remarkably illustrated by an experiment in which the leaves of the Agave americana, after twenty-four hours' exposure in the dark, were found to have absorbed only 0·3 of their volume of oxygen, while those of the fir, in which volatile oil is abundant, had taken up twice, and those of the oak, containing tannin, eighteen times as much oxygen.

In the flowers, both by day and night, there is a constant absorption of oxygen, and evolution of carbonic acid. In fact, an active oxidation is going on, attended by the evolution of heat, which, in the Arum maculatum and some other plants, is so great as to raise the temperature of the flower 10° or 12° above that of the surrounding air.

Decomposition of Water in the Plant.—In addition to the function which water performs in the plant, as the solvent of the different substances which form its nutriment, and hence as the medium through which they pass into its organs, it serves also as a direct food, undergoing decomposition, and yielding hydrogen to the organic substances. Its constituents, along with those of the carbonic acid absorbed, undergo a variety of transformations, and form the principal part of the non-nitrogenous constituents. It has been already observed that starch, sugar, and the other allied substances, may be considered as compounds of carbon with water; and they might be supposed to owe their origin to the carbonic acid losing the whole of its oxygen, and direct combination then ensuing between the residual carbon and a certain proportion of water; but this would imply that the latter substance undergoes no decomposition, and though undoubtedly the simplest view of the case, it is by no means the most probable. It is much more likely that the carbonic acid is only partially decomposed, half its oxygen being separated, and replaced by hydrogen, produced by the decomposition of a certain quantity of water into its elements. Thus, for instance, sugar may be produced from twelve equivalents of carbonic acid and twelve equivalents of water, twenty-four equivalents of oxygen being eliminated, as thus represented:

12equivalents ofcarbonic acid,C12O12O12
12"water,H12O12
1"sugar, and 24 of ox.C12H12O12 + O24

It must not be supposed that we are in a condition to assert that sugar is really produced in the manner here shown, the illustration being given merely for the purpose of pointing out how it may be supposed to occur, and on a similar principle it is possible to explain the formation of most other vegetable compounds; and this subject has been very fully discussed by the late Dr. Gregory, in his "Handbook of Organic Chemistry." That water must be decomposed, is evident from the fact, established by analysis, that the hydrogen of the plant generally exceeds the quantity required to form water with its oxygen, so that this excess at least must be produced by the decomposition of water. The hydrogen of the volatile oils, many of which contain no oxygen, and that of the fats, which contain only a small quantity, must manifestly be obtained in a similar manner.