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Advantages of AI over natural breeding

• Allows the choice of using the best possible bulls of proven quality in improving
the genetic make-up of the cattle population. Farmers have access to genes from
bulls of a quality which they may not individually afford. Frozen semen can be
transported globally.
• Disease control. Many potentially devastating diseases are spread by sexual
contact. Because of the extremely tight controls exerted over both the health of
donor bulls and the technical procedures themselves, these risks are vastly
reduced.
• Cost effectiveness. The cost of an AI straw is around £10, this is as nothing
compared with the costs of a Holstein bull (possibly £10,000 to buy). A bull is
expensive to rear, is relatively unproductive, vulnerable to disease or accident and
may even prove to be infertile.
• Flexibility. For a variety of reasons, a herdsman may not wish all calves to be
sired by a single bull with the same characteristics. It may well be impracticable
to keep sufficient bulls to cover all possible requirements.
• Safety. Although there are differences between breeds, any bull can be aggressive
and is potentially dangerous. This was a major stimulus to the initial setting up of
AI services.

Students should be aware of the principle of progeny testing in order to prove the quality
of a bull. They should know that:

• semen can be collected from bulls over a period of years and stored.
• samples of stored semen can be used to inseminate high quality cows and the
performance of the progeny can be measured. In this way, the quality of semen
from individual bulls is assessed before it is made available on a wide scale.
• the semen from quality bulls is used for large scale Al. Bulls of lesser quality are
culled. A bull can sire offspring over a period long exceeding its natural life.
Sometimes, when a bull has donated sufficient semen, it is culled to avoid
maintenance costs.

Flow chart illustrating progeny testing programme

1-2 year old bull provides semen

About 500 cows inseminated

9 months gestation

Calves have own offspring at 2½ years old; begin lactation

1 year of lactation

to kill pathogens. a buffer (usually citrate) to prevent pH changes due to. Dilution and storage of semen Semen is diluted in an 'extender'. milk or egg yolk to protect against cold shock (the initial cooling below body temperature). Henderson and Card had obtained in the use of laevulose solutions to protect fowl spermatozoa against the effects of freezing and thawing. Typically. This provides an appropriate concentration of spermatozoa. he will be up to 7 years old. 5. antibiotics. Performance result of calves Bull possibly in use for AI Thus. Some months later work resumed with the same material and negative results . lactic acid produced during sperm metabolism. as well as the correct overall water potential for their survival. the extender contains: 1. A dilution of around 50 times is usual. by the time the performance results of a bull's offspring are known. The extender also nourishes and protects the spermatozoa during storage and distribution. During this time. cryopreservation of spermatozoa remained an enigma until the serendipitous discovery of glycerol in 1948: "In the autumn of 1948 my colleagues Dr Audrey Smith and Dr C Polge were attempting to repeat the results which Shaffner. 4. allowing more inseminations from each sample. glucose (and/or other sugars) to provide an energy source for the spermatozoa. glycerol as a cryoprotectant (to protect damage due to the formation otherwise of ice crystals during freezing). 2. Small success attended the efforts and pending inspiration a number of solutions were put away in the cold store. Although it was known as early as 1776 that cooling semen held them in a dormant state and there followed a huge interest in the physics of cold. for example. 3. a great deal of semen can have been collected and stored. Equine Introduction.

however. whereas most cells are rendered incapable of fertilising an egg if frozen in the absence of a cryoprotectant. Tests with new material very soon showed that the albumen played no part in the protective effect. This very curious result suggested that chemical changes in the laevulose possibly caused or assisted by the flourishing growth of mould which had taken place during storage had produced a substance with surprising powers of protecting living cells against the effect of freezing and thawing. Obviously there had been some confusion with the various solutions although we never found out exactly what had happened. At this point. These pass through the sperm membrane and act both . which almost completely preserved motility in fowl spermatozoa frozen to -790C. He reported that the solution contained glycerol.were again obtained with all of the solutions except one. Since the discovery of glycerol protecting cells during freezing and thawing. fertilizing power. to some extent. the small amount (10-15 ml) of the miraculous solution remaining was handed over to our colleague Dr D Elliott for chemical analysis. water and a fair amount of protein. with some trepidation. Cryoprotectants A cryoprotectant such as glycerol allows a substantial percentage of spermatozoa in a sample to survive the freeze/thaw process and retain fertilising capability. Meanwhile further biological results had shown that not only was motility preserved after freezing and thawing but. progress has been dramatic in the area of multiple species gamete preservation. showed that the mysterious solution not only contained no unusual sugars but in fact contained no sugar at all.had been used in the course of morphological work on the spermatozoa at the same time as the laevulose solutions were being tested and with them had been put away in the cold store. also. Tests however. There are two general classes of cryoprotectants: 1) Penetrating cryoprotectants. For example the number of bull spermatozoa killed or rendered immotile during the freeze/thaw process has changed very little since 1955. there is still room for considerable improvement in the efficiency of the techniques. and our low temperature work became concentrated on the effects of glycerol in protecting living cells against the effects of low temperature (Parkes 1956).the glycerol and albumen of the histologist. It was then realised that Mayer’s albumen.

All solutes or colloids in a solution. Non-penetrating cryoprotectants move water out of the sperm which results in dehydration and shrinkage. Compounds placed into a solvent.intracellularly and extracellularly. from an environment rich in glycerol. lactose. Glycerol serves as a solute within the water and also penetrates into the sperm. and 2) Non-penetrating cryoprotectants. For example. All non-penetrating cryoprotectants. Despite this it is though that the primary beneficial effect of glycerol is extracellular. Most penetrating cryoprotectants serve as both a solvent and a solute. including proteins lipids and sugars are solutes or colloids and can not serve as a solvent. raffinose or trehalose. The higher the concentration of solutes the greater the osmotic pressure. Either in the extender or inside the spermatozoan. such as water. Non-penetrating cryprotectants include proteins such as in milk or egg yolk. The sperm reside in these unfrozen channels. A non-penetrating cryoprotectant such as lactose is a solute but because of the nature of the molecule it does not penetrate through the plasma membrane of a living cell. sugars such as fructose. These act only extracellularly. The process of thawing and insemination is potentially the most damaging to sperm due to rapid changes in volume. either within a spermatozoan or in the extracellular media. synthetic polymers such as polyvinylpyrrolidone or methyl cellulose and amides. remaining solvent plus solute. which dissolve and form a true solution or ionise are termed solutes. The movement of water out of the sperm is probably good because it is a direct result of water loss thus the possibility of intracellular ice damage is reduced. Thus lactose can contribute to the osmotic pressure of the seminal extender but not that of the sperm. lactose and salts of the extended semen with a high osmotic pressure (>1000 mOsmol/kg) to an environment in which the seminal . As extended semen is frozen crystals of pure water freeze into small blocks of ice between which are "channels" of unfrozen extender. contribute to the osmotic properties of the solution. Glycerol is the most common cryprotectant although DMSO and propylene glycol gave been used with some cells. Solvent cryoprotectants such as glycerol are beneficial because they function as a solvent with a freezing point much lower than that of water. glycerol contributes to the osmotic pressure of the extender or cell. mannose.

including the concentration of glycerol. The solution to the problem of rapid volume increase is to serially dilute the glycerol in the media surrounding the sperm in a controlled manner prior to insemination (similar to frozen/thawed embryos). However at least part of the problem of reduced fertility with cryopreserved sperm is a consequence of the apparent necessity to use a cryoprotectant such as glycerol. but optimal survival after re-warming. The ideal cooling and warming rates are also influenced by the composition of the extender. even when sperm are not subjected to the additional rigours of freezing and thawing. The outward movement of water is greater when sperm are cooled below 00C at a slow cooling rate than at a very rapid rate. There is abundant evidence that glycerol is toxic to sperm (Pace and Sullivan 1975). Typically straws of stallion sperm are warmed at 7000C/min by immersion in 370C water for ³ 30 seconds or 40000C/min by immersion in water at 750C for 7 seconds. the more rapid the cooling (freezing) rate. the more rapid should be the warming (thawing) rate.extender is diluted in the uterus (» 300 mOsmol/kg). Fortunately this is not necessary for most stallions spermatozoa as it would add another complex step to what needs to be a relatively simple procedure (to gain universal equine practitioner acceptance). results in a rapid increase in volume for the sperm as water moves in to equilibrate with the high concentration of intracellular glycerol (left behind). . Rapid swelling of the sperm may rupture the plasma membrane. In the event that a cooling rate is slower or faster than ideal the damage can be mitigated by appropriate adjustment of the warming rate. Ideally the cooling rate results in sufficient but not excessive outward movement of water and perhaps the formation of a few non- lethal ice crystals within each sperm. As a generalisation. The thawing procedure recommended by personnel who froze the spermatozoa should be followed closely.

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