It is only within the past 150 years that the sperm’s role in fertilization has been known. Anton van Leeuwenhoek, the Dutch microscopist who co-discovered sperm in the 1670s, first believed them to be parasitic animals living within the semen (hence the term spermatozoa, meaning “seed animals”). Although he originally assumed that they had nothing to do with reproducing the organism in which they were found, he later came to believe that each sperm contained a preformed embryo. Leeuwenhoek (1685) wrote that sperm were seeds (both sperma and semen mean “seed”) and that the female merely provided the nutrient soil in which the seeds were planted. In this, he was returning to a notion of procreation promulgated by Aristotle 2000 years earlier.
Try as he might, Leeuwenhoek was continually disappointed in his attempts to find preformed embryos within spermatozoa. Nicolas Hartsoeker, the other co-discoverer of sperm, drew a picture of what he hoped to find: a miniscule human (“homunculus”) within the sperm (Figure 4.1). This belief that the sperm contained the entire embryonic organism never gained much acceptance, as it implied an enormous waste of potential life. Most investigators regarded the sperm as unimportant.
The first evidence suggesting the importance of sperm in reproduction came from a series of experiments performed by Lazzaro Spallanzani in the late 1700s. Spallanzani induced male toads to ejaculate into taffeta breeches and found toad semen so filtered to be devoid of sperm; such semen did not fertilize eggs. He even showed that semen had to touch the eggs in order to be functional. However, Spallanzani (like many others) felt that the spermatic “animals” were parasites in the fluid; he thought the embryo was contained within the egg and needed spermatic fluid to activate it (see Pinto-Correia 1997).
The combination of better microscopic lenses and the elucidation of the cell theory (i.e., that all life is cellular, and all cells come from preexisting cells) led to a new appreciation of sperm function. In 1824, J. L. Prevost and J. B. Dumas claimed that sperm were not parasites, but rather the active agents of fertilization. They noted the universal existence of sperm in sexually mature males and their absence in immature and aged individuals. These observations, coupled with the known absence of sperm in the sterile mule, convinced Prevost and Dumas that “there exists an intimate relation between their presence in the organs and the fecundating capacity of the animal.” They proposed that the sperm entered the egg and contributed materially to the next generation.
These claims were largely disregarded until the 1840s, when A. von Kolliker described the formation of sperm from cells in the adult testes. He ridiculed the idea that the semen could be normal and yet support such an enormous number of parasites. Even so, von Kolliker denied there was any physical contact between sperm and egg. He believed that the sperm excited the egg to develop in much the same way a magnet communicates its presence to iron. It was not until 1876 that Oscar Hertwig and Herman Fol independently demonstrated sperm entry into the egg and the union of the two cells’ nuclei. Hertwig had been seeking an organism suitable for detailed microscopic observations, and he found the Mediterranean sea urchin (Paracentrotus lividus) to be perfect for this purpose. Not only was it common throughout the region and sexually mature throughout most of the year, but its eggs were available in large numbers and were transparent even at high magnifications.
1. Indeed, sperm was discovered around 1676, whereas the events of fertilization were not elucidated until 1876. Thus, for some 200 years people had no idea what the sperm actually did. See Pinto-Correia 1997 for details of this remarkable story.