As a general rule in small and medium-sized seeds the cuticle is well developed and represents the impermeable part of the seed coat, while in the cases of large seeds, such as those of _Adansonia gregorii_, _Mucuna gigantea_, _Wistaria maideniana_, and _Guilandina bonducella_, the cuticle is relatively unimportant and inconspicuous. In these seeds the extreme resistance which they exhibit appears to be located in the palisade cells.

In discussing the seed coat of _Melilotus alba_, Rees (33, p. 404) states that the outer layer consists of palisade cells covered, externally by a structureless membrane, which, however, did not appear to be cuticle but hemicellulose, as it stained magenta with chloriodid of zinc. The greater part of the walls of the palisade cells also appears to be composed of hemicellulose and the outer ends only were cuticularized. In order to find whether the outer membrane was in itself impermeable to water, this author treated seeds for short intervals in sulphuric acid to dissolve the outside covering without directly affecting the palisade cells. Seeds treated in this manner swelled in water and microscopic examination showed that the ends of the palisade cells were quite intact, but had separated from each other. From this it was concluded that the outer membrane is instrumental in conferring impermeability on the seed, although not directly responsible for it, as is the case with a true cuticle. It is further believed that it probably served as a cement substance by means of which the cuticularized ends of the cells were held together closely, thus forming a barrier through which water could not penetrate, but that as soon as this barrier was removed the ends of the palisade cells separated and water passed in between them.

USDA Bulletin No. 844 — Part 30

More than 20 years ago machines were devised by Kuntze, Michalowski (27, p. 86), Huss (15), and later by Hughes (14), to scarify impermeable seeds. Other methods have been recommended and employed to some extent for hastening the germination of seeds. Hiltner (13, p. 44) treated seeds of red clover, white clover, and alfalfa 10, 30, and 60 minutes with concentrated sulphuric acid and found that the best germination resulted from the 60-minute treatment. Love and Leighty (21) also treated the seeds of various legumes with concentrated sulphuric acid and obtained a better germination in all cases. In their investigations with _Melilotus alba_ it was found that a 2-hour treatment resulted in some injury to the seed, but that a treatment varying from 25 minutes to 1 hour gave good results. In most cases in our investigations the seed coats of sweet clover became permeable to water after a treatment of 15 minutes in concentrated sulphuric acid, and within 5 minutes all of the Malpighian cells were destroyed down to the light line. Harrington (10) found that the soil, season, climate, color, or size of red-clover seeds had no influence upon the percentage of impermeable seeds and that the good germination ordinarily obtained with red clover was due to the scarifying of the seed coats by the rasps of hulling machines. Harrington (11) also studied the agricultural value of impermeable seeds and found that alternations of temperature cause the softening and germinating of many impermeable clover seeds when a temperature of 10° C. or cooler is used in alternation with a temperature of 20° C. or warmer and that the effect of such an alternation of temperature is greatly increased by previously exposing the seeds to germinating conditions at a temperature of 10° C. or cooler and is decreased by previously exposing the seeds to germinating conditions at a temperature of 30° C. It is a well-known fact that impermeable seeds which remain in the field over winter germinate readily the following spring.

USDA Bulletin No. 844 — Part 30