Fertilization

Sexual reproduction starts with the combination of a sperm and an egg in a process called fertilization. This can occur either inside (internal fertilization) or outside (external fertilization) the body of the female. Humans provide an example of the former whereas seahorse reproduction is an example of the latter.

Fertilization, pictured in Figure 1a is the process in which gametes (an egg and sperm) fuse to form a zygote. The egg and sperm each contain one set of chromosomes. To ensure that the offspring has only one complete diploid set of chromosomes, only one sperm must fuse with one egg. In mammals, the egg is protected by a layer of extracellular matrix consisting mainly of glycoproteins called the zona pellucida. When a sperm binds to the zona pellucida, a series of biochemical events, called the acrosomal reactions, take place. In placental mammals, the acrosome contains digestive enzymes that initiate the degradation of the glycoprotein matrix protecting the egg and allowing the sperm plasma membrane to fuse with the egg plasma membrane, as illustrated in Figure 1b. The fusion of these two membranes creates an opening through which the sperm nucleus is transferred into the ovum. The nuclear membranes of the egg and sperm break down and the two haploid genomes condense to form a diploid genome.

Part A is a micrograph that shows a sperm whose head is touching the surface of an egg. The egg is much larger than the sperm. Part B is an illustration that shows the surface of the egg, which is coated with a zona pellucida. The sperm penetrates the zona pellucida and releases its DNA into the egg. At this point, changes in proteins just inside the egg’s cell membrane occur, preventing entry of other sperm.

Figure 1. (a) Fertilization is the process in which sperm and egg fuse to form a zygote. (b) Acrosomal reactions help the sperm degrade the glycoprotein matrix protecting the egg and allow the sperm to transfer its nucleus. (credit: (b) modification of work by Mariana Ruiz Villareal; scale-bar data from Matt Russell)

To ensure that no more than one sperm fertilizes the egg, once the acrosomal reactions take place at one location of the egg membrane, the egg releases proteins in other locations to prevent other sperm from fusing with the egg. If this mechanism fails, multiple sperm can fuse with the egg, resulting in polyspermy. The resulting embryo is not genetically viable and dies within a few days.

External Fertilization

External fertilization usually occurs in aquatic environments where both eggs and sperm are released into the water. After the sperm reaches the egg, fertilization takes place. Most external fertilization happens during the process of spawning where one or several females release their eggs and the male(s) release sperm in the same area, at the same time. The release of the reproductive material may be triggered by water temperature or the length of daylight. Nearly all fish spawn, as do crustaceans (such as crabs and shrimp), mollusks (such as oysters), squid, and echinoderms (such as sea urchins and sea cucumbers). Figure 2a shows salmon spawning in a shallow stream. Frogs, like those shown in Figure 2b, corals, mayflies, and mosquitoes also spawn.

Photo A shows many salmon swimming up a shallow creek. Photo B shows mating toads.

Figure 2. (a) Salmon reproduce through spawning. (b) During sexual reproduction in toads, the male grasps the female from behind and externally fertilizes the eggs as they are deposited. (credit a: Dan Bennett; credit b: “OakleyOriginals”/Flickr)

Pairs of fish that are not broadcast spawners may exhibit courtship behavior. This allows the female to select a particular male. The trigger for egg and sperm release (spawning) causes the egg and sperm to be placed in a small area, enhancing the possibility of fertilization.

External fertilization in an aquatic environment protects the eggs from drying out. Broadcast spawning can result in a greater mixture of the genes within a group, leading to higher genetic diversity and a greater chance of species survival in a hostile environment. For sessile aquatic organisms like sponges, broadcast spawning is the only mechanism for fertilization and colonization of new environments. The presence of the fertilized eggs and developing young in the water provides opportunities for predation resulting in a loss of offspring. Therefore, millions of eggs must be produced by individuals, and the offspring produced through this method must mature rapidly. The survival rate of eggs produced through broadcast spawning is low.

Internal Fertilization

Internal fertilization occurs most often in land-based animals, although some aquatic animals also use this method. There are three ways that offspring are produced following internal fertilization. In oviparity, fertilized eggs are laid outside the female’s body and develop there, receiving nourishment from the yolk that is a part of the egg. This occurs in most bony fish, many reptiles, some cartilaginous fish, most amphibians, two mammals, and all birds. Reptiles and insects produce leathery eggs, while birds and turtles produce eggs with high concentrations of calcium carbonate in the shell, making them hard. Chicken eggs are an example of this second type.

In ovoviparity, fertilized eggs are retained in the female, but the embryo obtains its nourishment from the egg’s yolk and the young are fully developed when they are hatched. This occurs in some bony fish (like the guppy Lebistes reticulatus), some sharks, some lizards, some snakes (such as the garter snake Thamnophis sirtalis), some vipers, and some invertebrate animals (like the Madagascar hissing cockroach Gromphadorhina portentosa).

In viviparity the young develop within the female, receiving nourishment from the mother’s blood through a placenta. The offspring develops in the female and is born alive. This occurs in most mammals, some cartilaginous fish, and a few reptiles.

Internal fertilization has the advantage of protecting the fertilized egg from dehydration on land. The embryo is isolated within the female, which limits predation on the young. Internal fertilization enhances the fertilization of eggs by a specific male. Fewer offspring are produced through this method, but their survival rate is higher than that for external fertilization.

The Evolution of Reproduction

Once multicellular organisms evolved and developed specialized cells, some also developed tissues and organs with specialized functions. An early development in reproduction occurred in the Annelids. These organisms produce sperm and eggs from undifferentiated cells in their coelom and store them in that cavity. When the coelom becomes filled, the cells are released through an excretory opening or by the body splitting open. Reproductive organs evolved with the development of gonads that produce sperm and eggs. These cells went through meiosis, an adaption of mitosis, which reduced the number of chromosomes in each reproductive cell by half, while increasing the number of cells through cell division.

Complete reproductive systems were developed in insects, with separate sexes. Sperm are made in testes and then travel through coiled tubes to the epididymis for storage. Eggs mature in the ovary. When they are released from the ovary, they travel to the uterine tubes for fertilization. Some insects have a specialized sac, called a spermatheca, which stores sperm for later use, sometimes up to a year. Fertilization can be timed with environmental or food conditions that are optimal for offspring survival.

Vertebrates have similar structures, with a few differences. Non-mammals, such as birds and reptiles, have a common body opening, called a cloaca, for the digestive, excretory and reproductive systems. Coupling between birds usually involves positioning the cloaca openings opposite each other for transfer of sperm. Mammals have separate openings for the systems in the female and a uterus for support of developing offspring. The uterus has two chambers in species that produce large numbers of offspring at a time, while species that produce one offspring, such as primates, have a single uterus.

Sperm transfer from the male to the female during reproduction ranges from releasing the sperm into the watery environment for external fertilization, to the joining of cloaca in birds, to the development of a penis for direct delivery into the female’s vagina in mammals.

In Summary: Fertilization

Sexual reproduction starts with the combination of a sperm and an egg in a process called fertilization. This can occur either outside the bodies or inside the female. Both methods have advantages and disadvantages. Once fertilized, the eggs can develop inside the female or outside. If the egg develops outside the body, it usually has a protective covering over it. Animal anatomy evolved various ways to fertilize, hold, or expel the egg. The method of fertilization varies among animals. Some species release the egg and sperm into the environment, some species retain the egg and receive the sperm into the female body and then expel the developing embryo covered with shell, while still other species retain the developing offspring through the gestation period.