ANGLEINTENSITY
901.00
80.98
70.94
60.87
50.77
40.64
30.50
20.34
10.17

In the sunlight, it makes no difference at what height a light reading is taken. In forest and thicket as well as in some herbaceous formations, the intensity of the light, if there is any difference, is greatest just beneath the foliage of the facies. In forests especially, the light is increasingly diffuse toward the ground, particularly where layers intervene. In woodland formations, moreover, the exact spot in which a reading is made must be carefully chosen, unless the foliage is so dense that the shade is uniform. A very satisfactory plan is to take readings in two or more spots where the shade appears to be typical, and to make a check reading in a “sunfleck,” a spot where sunlight shows through. In forests and thickets, the sunflecks are fleeting, and the light value is practically that of the shade. In passing into open woodland and thicket, the sunflecks increase in size and permanence, until finally they exceed the shade areas in amount and become typical of the formation.

Reflected and Absorbed Light

88. The fate of incident light. The light present in a habitat and incident upon a leaf is not all available for photosynthesis. Part is reflected or screened out by the epidermis, and a certain amount passes through the chlorenchym, except in very thick leaves. The light absorbed is by far the greatest in the majority of species. Many plants with dense coatings of hairs reflect or withhold more light than they absorb, and the amount of light reflected by a thick cuticule is likewise great. As light is imponderable, the actual amount absorbed or reflected by the leaf can not be determined. It is possible, however, to express this in terms of the total amount received, by means of readings with solio paper, and the knowledge thus obtained is of great importance in interpreting the modifications of certain types of leaves. For example, a leaf with a densely hairy epidermis may receive light of the full intensity, 1; the amount reflected or screened out by the hairs may be 95 per cent of this, the amount absorbed 5 per cent, and that transmitted, nil. In the majority of cases, however, the absorbed light is considerably more than the amount reflected or transmitted.

Fig. 14. Leaf print: exposed 10 m., 11 A.M. August 20. The leaves are from sun and shade forms of Bursa bursa-pastoris, Rosa sayii, Thalictrum sparsiflorum, and Machaeranthera aspera. In each the shade leaf prints more deeply.

89. Methods of determination. If results are to be of value, reflected and transmitted light must be determined in the habitat of the plant simultaneously with the total light which a leaf receives. An approximation of the light reflected from a leaf surface is secured by placing the photometer so that the light reflected is thrown upon the solio strip. A much more satisfactory method, however, is to determine it in connection with the amount of light transmitted through the epidermis. This is done by stripping a piece of epidermis from the upper surface of the leaf and placing it over the slit in the photometer for an exposure. An exposure in the full light of the habitat is made simultaneously with another photometer, or immediately afterward upon the same strip. When the epidermis is not too dense, both exposures are permitted to reach the same tint, and the relation between them is precisely that of their lengths of exposure. Ordinarily the two exposures are made absolutely simultaneous by placing the epidermis over half of the opening, leaving the other half to record the full light value, and the results, or epidermis prints, are referred to a multiple standard. The difference between the two values thus obtained represents the amount of reflected light together with that screened by the epidermis. The amount of light transmitted through the leaf may be measured in the same way by using the leaf itself in place of the epidermis alone. The time of exposure is necessarily long, however, and it has been found practicable to obtain leaf prints by exposing the leaf in a printing frame, upon solio paper, at the same time that the epidermis print is made. In a few species both the upper and lower epidermis can be removed and the amount of light absorbed determined directly by exposing the strip covered with the chlorenchym. Generally, however, this must be computed by subtracting the sum of the per cents of reflected and transmitted light from 100 per cent, which represents the total light.

Fig. 15. Leaf print: exposure as before. Sun and shade leaves of Achillea lanulosa, Capnoides aureum, Antennaria umbrinella, Galium boreale, and Potentilla propinqua.

90. Leaf and epidermis prints. In diphotic leaves the screening effect of the lower epidermis may be ignored. Isophotic sun leaves, i. e., those nearly upright in position or found above light-colored, reflecting soils, are usually strongly illuminated on both sides, and the absorbed light can be obtained only by measuring the screening effect of both epiderms. Shade leaves and submerged leaves often contain chloroplasts in the epidermis, and the above method can not be applied to them. In fact, in habitats where the light is quite diffuse, practically all incident light is absorbed. The rare exceptions are those shade leaves with a distinct bloom. In addition to their use in obtaining the amount of light absorbed, both leaf and epidermis prints are extremely interesting for the direct comparison of light relations in the leaves of species belonging to different habitats. The relative screening value of the upper and lower epidermis, or of the corresponding epiderms of two ecads or two species, is readily ascertained by exposing the two side by side in sunshine, over the slit in the photometer. For leaf prints fresh leaves are desirable, though nearly the same results can be obtained from leaves dried under pressure. The leaves are grouped as desired on the glass of a printing frame, and covered with a sheet of solio. They are then exposed to full sunlight, preferably at meridian, and the prints evaluated by means of the multiple standard. This method is especially useful in the comparison of ecads of one species. These differences due to transmitted light are very graphic, and can easily be preserved by “toning” the print in the usual way.