His most valuable work in this direction is a paper on a genus Kingia, discovered in New Holland in 1825; the structure of the seeds in this genus led him to seek more accurate knowledge of the unfertilised ovule in the Phanerogams generally, and especially in the Cycads and Conifers. In spite of the labours of Gärtner and the later researches of Treviranus, there was still considerable obscurity attaching to the theory of the seed, for no one had yet succeeded in referring the position of the embryo in the ripe seed to a general law. For this it was necessary to submit the ovule before fertilisation to careful examination, and Robert Brown carried out this first step to a history of development with great success; he was the first to distinguish the integuments and the nucleus in the ovule, and the embryo-sac in the nucleus, parts which Malpighi and Grew had indeed observed but had not brought out with perfect clearness. The micropyle and the hilum of the seed had not yet been properly distinguished, but had been to some extent even confounded with one another. Robert Brown showed that the hilum answers to the point of attachment of the ovule, while the micropyle is a canal formed by the integuments of the ovule and leading to the summit of the nucleus; that in anatropous ovules the micropyle lies beside the hilum, in orthotropous ovules opposite to it; that the embryo in the embryo-sac (amnion) is always formed at the spot which lies nearest the micropyle, and that the radicle of the embryo is always turned towards the micropyle,—facts which at once established the general rule by which to determine the position of the embryo in the seed and in the fruit. He also gave the first correct explanation of the endosperm as a nourishing substance formed inside the embryo-sac after fertilisation, and more than this, he was the first to distinguish the perisperm as a substance formed outside the embryo-sac in the tissue of the nucleus.
In this way Robert Brown established morphological relations in the organisation of the seed of the Monocotyledons and Dicotyledons, which count among the most important
