Difference between revisions of "Oestrus Behaviour - Anatomy & Physiology"
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− | + | == Introduction == | |
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− | = Introduction = | ||
Oestrus is the period of sexual receptivity in a female, commonly referred to as 'heat'. | Oestrus is the period of sexual receptivity in a female, commonly referred to as 'heat'. | ||
− | For further infomation on oestrous cyclicity, follow [[ | + | For further infomation on oestrous cyclicity, follow [[Oestrous_Cycle_- Anatomy & Physiology|this link]]. |
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---- | ---- | ||
− | = Cow = | + | ==Cow== |
[[Image:Oestrus Behaviour cow.jpg|thumb|right|150px|<p>Expression of Oestrus Behaviour in Relation to Ovulation</p><sup> Courtesy of In Practice,November 2005.</sup>]] | [[Image:Oestrus Behaviour cow.jpg|thumb|right|150px|<p>Expression of Oestrus Behaviour in Relation to Ovulation</p><sup> Courtesy of In Practice,November 2005.</sup>]] | ||
− | == Oestrus Detection == | + | ===Oestrus Detection=== |
* Oestrus detection is critical, as the absence of detection is the most important reason for delayed breeding. | * Oestrus detection is critical, as the absence of detection is the most important reason for delayed breeding. | ||
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* Average oestrus detection rates are ~ 60%, the target is >70%! | * Average oestrus detection rates are ~ 60%, the target is >70%! | ||
− | === Standing to be Mounted === | + | ==== Standing to be Mounted ==== |
Although pedometers are increasingly being used and in-line milk progesterone tests are being developed, the most reliable method for determining the optimum timing of artificial insemination (AI) is the observation of cows standing to be mounted (STBM). | Although pedometers are increasingly being used and in-line milk progesterone tests are being developed, the most reliable method for determining the optimum timing of artificial insemination (AI) is the observation of cows standing to be mounted (STBM). | ||
* Ovulation occurs approximately 30 hours after the first display of STBM, which in turn is 24 hours after the LH surge. | * Ovulation occurs approximately 30 hours after the first display of STBM, which in turn is 24 hours after the LH surge. | ||
* The optimum time to inseminate in order to achieve the best conception rates is six to 24 hours after the first observed STBM. | * The optimum time to inseminate in order to achieve the best conception rates is six to 24 hours after the first observed STBM. | ||
− | == Signs of Oestrus == | + | === Signs of Oestrus === |
* Standing to be mounted is the only truly reliable indication of oestrus. | * Standing to be mounted is the only truly reliable indication of oestrus. | ||
* Other signs of oestrus may include: | * Other signs of oestrus may include: | ||
** Homosexual mounting | ** Homosexual mounting | ||
− | ** Bellowing when | + | ** Bellowing when isolated |
** Holding milk | ** Holding milk | ||
** Brawling | ** Brawling | ||
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** Mucus discharge (also when coming into and going out of oestrus). This is called a '''bulling string'''. | ** Mucus discharge (also when coming into and going out of oestrus). This is called a '''bulling string'''. | ||
− | === Factors Influencing Oestrus Behaviour === | + | ==== Factors Influencing Oestrus Behaviour ==== |
Sexual behaviour in cows is itself influenced by numerous factors: | Sexual behaviour in cows is itself influenced by numerous factors: | ||
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** Cow–cow interactions are reduced, thus interfering with oestrus detection. | ** Cow–cow interactions are reduced, thus interfering with oestrus detection. | ||
− | == Aids to Improve Oestrus Detection == | + | === Aids to Improve Oestrus Detection === |
− | For methods of oestrous synchronisation and oestrus induction click [[ | + | For methods of oestrous synchronisation and oestrus induction click [[Oestrous Cycle Pharmacological Manipulation - Anatomy & Physiology|here]]. |
− | === Tail Paint === | + | ==== Tail Paint ==== |
* Applied to the base of the tail and sacrum. | * Applied to the base of the tail and sacrum. | ||
Line 85: | Line 79: | ||
* Cheap and effective | * Cheap and effective | ||
− | === KaMaR Heat Mount Detectors === | + | ==== KaMaR Heat Mount Detectors ==== |
* More expensive | * More expensive | ||
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* AM/PM rule still holds: If you see a red detector in the morning, breed her late that afternoon. If you see a red detector in the evening, breed her the next morning. | * AM/PM rule still holds: If you see a red detector in the morning, breed her late that afternoon. If you see a red detector in the evening, breed her the next morning. | ||
− | === Closed Circuit Television (CCTV) === | + | ==== Closed Circuit Television (CCTV) ==== |
* CCTV with a time-lapsed video | * CCTV with a time-lapsed video | ||
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* Good cow identification is important. | * Good cow identification is important. | ||
− | === Teaser Bulls === | + | ==== Teaser Bulls ==== |
− | * Teaser bulls or androgenised cows will identify cows | + | * Teaser bulls or androgenised cows will identify cows that are in oestrus provided they have some sort of marker, such as a '''chin-ball device'''. |
** The halter has a cone shaped device underneath which contains a freely rotating ball, working on the same principle as a ball-pen. | ** The halter has a cone shaped device underneath which contains a freely rotating ball, working on the same principle as a ball-pen. | ||
** The stainless steel ball container can be filled with ink of the desired colour. | ** The stainless steel ball container can be filled with ink of the desired colour. | ||
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* Some bulls aquire harems of specific cows. | * Some bulls aquire harems of specific cows. | ||
− | == Clinical Methods and Assessments for Oestrus Detection == | + | === Clinical Methods and Assessments for Oestrus Detection === |
Successful oestrus detection is reliant both on observation and the display of oestrous | Successful oestrus detection is reliant both on observation and the display of oestrous | ||
behaviour. If either element is reduced, this has serious effects on detection and thus submission rates.A period of 30 minutes observation at least three times daily is recommended. | behaviour. If either element is reduced, this has serious effects on detection and thus submission rates.A period of 30 minutes observation at least three times daily is recommended. | ||
− | === Progesterone Levels === | + | ==== Progesterone Levels ==== |
* Low plasma or milk progesterone indicates that the cow is close to, or in oestrus. | * Low plasma or milk progesterone indicates that the cow is close to, or in oestrus. | ||
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* Sampling of milk progesterone to check oestrus identification can be counterproductive: aware that the purpose of this measurement is to ‘investigate’ their accuracy of oestrus detection, farm staff may be tempted only to sample cows that are definitely in oestrus during the period of checks. | * Sampling of milk progesterone to check oestrus identification can be counterproductive: aware that the purpose of this measurement is to ‘investigate’ their accuracy of oestrus detection, farm staff may be tempted only to sample cows that are definitely in oestrus during the period of checks. | ||
− | === Other Methods === | + | ==== Other Methods ==== |
− | * Detection of a [[ | + | * Detection of a [[Follicles - Anatomy & Physiology|'''follicle''']] and absence of a [[Corpus_Luteum_- Anatomy & Physiology|'''corpus luteum''']] by palpation or ultrasound examination of [[Ovary_- Anatomy & Physiology|ovaries]]. |
− | * Crystallisation patterns in [[ | + | * Crystallisation patterns in [[Vagina and Vestibule - Anatomy & Physiology|vaginal]] mucus. |
* Fall in body temperature prior to oestrus. | * Fall in body temperature prior to oestrus. | ||
− | * Reduction in [[ | + | * Reduction in [[Vagina and Vestibule - Anatomy & Physiology|vaginal]] pH. |
---- | ---- | ||
− | = Ewe = | + | == Ewe == |
− | == Oestrus Detection == | + | === Oestrus Detection === |
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− | == Signs of Oestrus == | + | === Signs of Oestrus === |
* Restless | * Restless | ||
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− | == Clinical Methods and Assessments for Oestrus Detection == | + | === Clinical Methods and Assessments for Oestrus Detection === |
* Low plasma progesterone | * Low plasma progesterone | ||
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---- | ---- | ||
− | = Sow = | + | == Sow == |
− | == Oestrus Detection == | + | === Oestrus Detection === |
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− | == Signs of Oestrus == | + | === Signs of Oestrus === |
− | * Stands for a boar producing lots of [[ | + | * Stands for a boar producing lots of [[Attractivity Behaviour - Anatomy & Physiology#Pheramones|pheromones]] in [[Salivary_Glands_-_Anatomy_&_Physiology#Saliva|saliva]]. |
* Stands very firmly to be mounted, hence the 'back pressure test' is very reliable. | * Stands very firmly to be mounted, hence the 'back pressure test' is very reliable. | ||
** But, becomes refractory after 10 minutes standing, then no visible response for 1-2 hours. | ** But, becomes refractory after 10 minutes standing, then no visible response for 1-2 hours. | ||
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*** Nosing | *** Nosing | ||
*** Mounting other females | *** Mounting other females | ||
− | ** Increase in [[ | + | ** Increase in [[Vagina and Vestibule - Anatomy & Physiology|vaginal]] mucous (requires digital examination) |
*** Usually more pronounced in guilts than sows. | *** Usually more pronounced in guilts than sows. | ||
---- | ---- | ||
− | = Mare = | + | == Mare == |
* Oestrous detection is a problem because mares not in oestrus can cause significant damage to a stallion. | * Oestrous detection is a problem because mares not in oestrus can cause significant damage to a stallion. | ||
− | * May be a slight [[ | + | * May be a slight [[Oedema|oedema]] of the vulva and a mucoid discharge. |
* Generally mares return to oestrous about 6-10 days after foaling. | * Generally mares return to oestrous about 6-10 days after foaling. | ||
− | ** Called a ''''foal heat'''' because often the foal develops [[ | + | ** Called a ''''foal heat'''' because often the foal develops [[Diarrhoea|diarrhoea]] at this time, which makes the oestrus easy to detect. |
*** Fertility may not be high at 'foal heat' oestrus. | *** Fertility may not be high at 'foal heat' oestrus. | ||
* Mares with an older foal at foot may not show signs of oestrus, as they are protective of the foal (''''foal shy''''). | * Mares with an older foal at foot may not show signs of oestrus, as they are protective of the foal (''''foal shy''''). | ||
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* Mare will stand as if to urinate, lift her tail and lean her hindquarters toward the stallion. | * Mare will stand as if to urinate, lift her tail and lean her hindquarters toward the stallion. | ||
* Mare repeatedly exposes her clitoris ('winking') | * Mare repeatedly exposes her clitoris ('winking') | ||
− | * If the mare is in oestrus, the stallion usually demonstrates a [[ | + | * If the mare is in oestrus, the stallion usually demonstrates a [[Pre-copulatory Behaviour - Anatomy & Physiology#Flehmen_Response|Flehmen response]]. |
− | == Clinical Methods and Assessments for Oestrus Detection == | + | === Clinical Methods and Assessments for Oestrus Detection === |
− | === Ultrasound === | + | ==== Ultrasound ==== |
− | * Detection of a [[ | + | * Detection of a [[Follicles - Anatomy & Physiology|'''follicle''']] and absence of a [[Corpus Luteum - Anatomy & Physiology|'''corpus luteum''']] by palpation or ultrasound examination of [[Ovary_- Anatomy & Physiology|ovaries]]. |
* Demonstration of follicle size, follicle wall thickening and follicle wall softening on ultrasound. | * Demonstration of follicle size, follicle wall thickening and follicle wall softening on ultrasound. | ||
− | === Vaginal and Cervical Examination === | + | ==== Vaginal and Cervical Examination ==== |
− | By visual; examination of the [[ | + | By visual; examination of the [[Vagina and Vestibule - Anatomy & Physiology|vagina]] and [[Cervix_- Anatomy & Physiology|cervix]] using an illuminated speculum, it is possible to detect the preovulation period. |
− | * In dioestrus, the [[ | + | * In dioestrus, the [[Cervix_- Anatomy & Physiology|cervix]] is small, constricted and firm. |
− | * In dioestrus, the [[ | + | * In dioestrus, the [[Cervix_- Anatomy & Physiology|cervix]] and [[Vagina and Vestibule - Anatomy & Physiology|vagina]] are pale pink and mucus is scanty and sticky. |
− | * During oestrus, there is a gradual increase in vascularity of the genital tract and relaxation of the [[ | + | * During oestrus, there is a gradual increase in vascularity of the genital tract and relaxation of the [[Cervix_- Anatomy & Physiology|cervix]] with dilation of the cervical os. |
− | * Approaching [[ | + | * Approaching [[Ovulation - Anatomy & Physiology|ovulation]], the [[Cervix_- Anatomy & Physiology|cervix]] becomes very relaxed and its protrusion can be seen on the [[Vagina and Vestibule - Anatomy & Physiology|vaginal]] floor. |
− | ** [[ | + | ** [[Cervix_- Anatomy & Physiology|Cervical]] folds are oedematous |
− | * [[ | + | * [[Vagina and Vestibule - Anatomy & Physiology|Vaginal]] folds glisten with clear lubricant mucus. |
---- | ---- | ||
− | = Bitch = | + | == Bitch == |
− | * Long period of '''proestrus''' where the bitch is [[ | + | * Long period of '''proestrus''' where the bitch is [[Attractivity Behaviour - Anatomy & Physiology|attractive]], but will not allow mating. |
** Characterized by: | ** Characterized by: | ||
*** Vulval swelling | *** Vulval swelling | ||
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** Deviation of the tail | ** Deviation of the tail | ||
** Discharge becomes paler in colour | ** Discharge becomes paler in colour | ||
− | ** Reduction in vulval [[ | + | ** Reduction in vulval [[Oedema|oedema]] |
− | * Poor relationship between time of [[ | + | * Poor relationship between time of [[Ovulation - Anatomy & Physiology|ovulation]] and external appearance. |
− | |||
− | |||
− | |||
− | |||
+ | === Clinical Methods and Assessments for Oestrus Detection === | ||
* Assessment of vulval softening | * Assessment of vulval softening | ||
− | * Assessment of [[ | + | * Assessment of [[Vagina and Vestibule - Anatomy & Physiology#Histology_and_Innervation|vaginal epithelial cells]] |
− | * Examination of the appearance of the [[ | + | * Examination of the appearance of the [[Vagina and Vestibule - Anatomy & Physiology|vaginal]] wall. |
− | * Measurement of plasma progesterone concentration (detecting a rise in progesterone that preceeds [[ | + | * Measurement of plasma progesterone concentration (detecting a rise in progesterone that preceeds [[Ovulation - Anatomy & Physiology|'''ovulation''']] in this species). |
---- | ---- | ||
− | = Queen = | + | == Queen == |
− | * Oestrus detection is important, for although the queen is an '''induced ovulator''' (thus [[ | + | * Oestrus detection is important, for although the queen is an '''induced ovulator''' (thus [[Ovulation - Anatomy & Physiology|'''ovulation''']] will not be missed), the LH response to mating decreases with each day of oestrus and [[Follicles - Anatomy & Physiology|'''follicles''']] ultimately regress. |
* '''Proestrus''' is short and often unnoticed. | * '''Proestrus''' is short and often unnoticed. | ||
− | * Oestrus is characterized by increased [[ | + | * Oestrus is characterized by increased [[Attractivity Behaviour - Anatomy & Physiology#Auditory_Signals|vocalization]],rubbing and rolling. |
* Queen is more active and will seek the tom. | * Queen is more active and will seek the tom. | ||
* Stands to be mated by lowering her front quarters and extending her hind legs. | * Stands to be mated by lowering her front quarters and extending her hind legs. | ||
− | * Demonstrates [[ | + | * Demonstrates [[Receptivity Behaviour - Anatomy & Physiology|lordosis]] |
* Tail erect and held to one side. | * Tail erect and held to one side. | ||
* Slight mucoid vulval discharge. | * Slight mucoid vulval discharge. | ||
− | + | {{Learning | |
− | < | + | |flashcards = [[Equine Reproduction and Stud Medicine Q&A 15]]<br>[[Equine Reproduction and Stud Medicine Q&A 16]] |
− | + | }} | |
− | + | [[Category:Reproductive Behaviour]] | |
+ | [[Category:Bullet Points]] |
Latest revision as of 13:44, 5 July 2012
Introduction
Oestrus is the period of sexual receptivity in a female, commonly referred to as 'heat'. For further infomation on oestrous cyclicity, follow this link.
Species | Mare | Cow | Ewe | Sow | Bitch |
---|---|---|---|---|---|
Duration of Oestrus | 7 days (2-12 days) | 15 hours (6-24 hours) | 30 hours (18-48 hours) | 50 hours (12-96 hours) | 9 days (4-21 days) |
Cow
Oestrus Detection
- Oestrus detection is critical, as the absence of detection is the most important reason for delayed breeding.
- A common problem is lack of return to oestrus at a reliable time after parturition.
- Cows are not often run with the bull, and so detection is the job of the herdsperson.
- 70% of oestrus activity occurs at night.
- Groups of cows need to be examined at least 4 times a day at equally spread intervals.
- Average oestrus detection rates are ~ 60%, the target is >70%!
Standing to be Mounted
Although pedometers are increasingly being used and in-line milk progesterone tests are being developed, the most reliable method for determining the optimum timing of artificial insemination (AI) is the observation of cows standing to be mounted (STBM).
- Ovulation occurs approximately 30 hours after the first display of STBM, which in turn is 24 hours after the LH surge.
- The optimum time to inseminate in order to achieve the best conception rates is six to 24 hours after the first observed STBM.
Signs of Oestrus
- Standing to be mounted is the only truly reliable indication of oestrus.
- Other signs of oestrus may include:
- Homosexual mounting
- Bellowing when isolated
- Holding milk
- Brawling
- Restlessness
- Chin pressing
- Swelling of vulva (also when coming into and going out of oestrus).
- Mucus discharge (also when coming into and going out of oestrus). This is called a bulling string.
Factors Influencing Oestrus Behaviour
Sexual behaviour in cows is itself influenced by numerous factors:
- Cows tend to be ridden by cows in, coming into or going out of oestrus. The more sexually active females there are, the greater the number of mounts and likelihood of cows being observed standing to be mounted.
- Lame cows are less likely to show signs of sexual behaviour.
- Non-slip surfaces encourage sexual activity, while slippery and rough surfaces discourage such behaviour.
- Lack of space is prohibitive.
- The presence of a stock bull can be counterproductive.
- Rarely see a cow actually being served
- Cow–cow interactions are reduced, thus interfering with oestrus detection.
Aids to Improve Oestrus Detection
For methods of oestrous synchronisation and oestrus induction click here.
Tail Paint
- Applied to the base of the tail and sacrum.
- Removed by rubbing when the cow stands to be ridden.
- Cheap and effective
KaMaR Heat Mount Detectors
- More expensive
- Must be identified, as they can become displaced.
- Pressure sensitive device with a built-in timing mechanism designed to be activated by standing heat behavior.
- Glued onto the sacrum
- Pressure from the brisket of a mounting animal requires approximately 3 seconds to turn the detector from white to red.
- This timing mechanism helps distinguish between true standing heat versus false mounting activity.
- AM/PM rule still holds: If you see a red detector in the morning, breed her late that afternoon. If you see a red detector in the evening, breed her the next morning.
Closed Circuit Television (CCTV)
- CCTV with a time-lapsed video
- Useful during night hours when the cows are not observed.
- Good cow identification is important.
- Teaser bulls or androgenised cows will identify cows that are in oestrus provided they have some sort of marker, such as a chin-ball device.
- The halter has a cone shaped device underneath which contains a freely rotating ball, working on the same principle as a ball-pen.
- The stainless steel ball container can be filled with ink of the desired colour.
- Each fill is enough to mark 25-30 cows (approx. 0.5 litre).
- Marks fade in 4 to 14 days
- Using bulls risks the spread of venereal disease.
- Some bulls aquire harems of specific cows.
Clinical Methods and Assessments for Oestrus Detection
Successful oestrus detection is reliant both on observation and the display of oestrous behaviour. If either element is reduced, this has serious effects on detection and thus submission rates.A period of 30 minutes observation at least three times daily is recommended.
Progesterone Levels
- Low plasma or milk progesterone indicates that the cow is close to, or in oestrus.
- Milk progesterone assays can be used routinely on alternate days from 25-30 days postpartum.
- Sampling of milk progesterone to check oestrus identification can be counterproductive: aware that the purpose of this measurement is to ‘investigate’ their accuracy of oestrus detection, farm staff may be tempted only to sample cows that are definitely in oestrus during the period of checks.
Other Methods
- Detection of a follicle and absence of a corpus luteum by palpation or ultrasound examination of ovaries.
- Crystallisation patterns in vaginal mucus.
- Fall in body temperature prior to oestrus.
- Reduction in vaginal pH.
Ewe
Oestrus Detection
- Oestrus detection is only important when Artificial Insemination regimes are practiced and often these are now done following hormonal stimulation rather than waiting for a spontaneous cycle.
Signs of Oestrus
- Restless
- Seeks ram
- Forms a following harem with other ewes
- Waggles tail
- Moves tail laterally to allow mating
- Slightly swollen vulva
- Slight mucoid vulval discharge
Clinical Methods and Assessments for Oestrus Detection
- Low plasma progesterone
Sow
Oestrus Detection
- Oestrus detection is a crucial assessment to ensure sows are presented to the boar at the correct time.
- Good relationship between weaning and return of oestrus, unlike the cow.
- Observe twice daily remote from feeding time.
Signs of Oestrus
- Stands for a boar producing lots of pheromones in saliva.
- Stands very firmly to be mounted, hence the 'back pressure test' is very reliable.
- But, becomes refractory after 10 minutes standing, then no visible response for 1-2 hours.
- Vulva swollen and congested 3 days prior to Oestrus.
- Desire to seek the boar.
- Other signs include:
- Restlessness with repetitive grunts
- Pacing
- Searching
- Mounting
- Being mounted
- Lowered feed intake
- Loss of appetite
- Male-like sexual behaviour:
- Persuing
- Nosing
- Mounting other females
- Increase in vaginal mucous (requires digital examination)
- Usually more pronounced in guilts than sows.
Mare
- Oestrous detection is a problem because mares not in oestrus can cause significant damage to a stallion.
- May be a slight oedema of the vulva and a mucoid discharge.
- Generally mares return to oestrous about 6-10 days after foaling.
- Called a 'foal heat' because often the foal develops diarrhoea at this time, which makes the oestrus easy to detect.
- Fertility may not be high at 'foal heat' oestrus.
- Called a 'foal heat' because often the foal develops diarrhoea at this time, which makes the oestrus easy to detect.
- Mares with an older foal at foot may not show signs of oestrus, as they are protective of the foal ('foal shy').
- Commonly teasing is performed (usually daily) using a less valuable stallion over a 'teasing board'.
- Mare will stand as if to urinate, lift her tail and lean her hindquarters toward the stallion.
- Mare repeatedly exposes her clitoris ('winking')
- If the mare is in oestrus, the stallion usually demonstrates a Flehmen response.
Clinical Methods and Assessments for Oestrus Detection
Ultrasound
- Detection of a follicle and absence of a corpus luteum by palpation or ultrasound examination of ovaries.
- Demonstration of follicle size, follicle wall thickening and follicle wall softening on ultrasound.
Vaginal and Cervical Examination
By visual; examination of the vagina and cervix using an illuminated speculum, it is possible to detect the preovulation period.
- In dioestrus, the cervix is small, constricted and firm.
- In dioestrus, the cervix and vagina are pale pink and mucus is scanty and sticky.
- During oestrus, there is a gradual increase in vascularity of the genital tract and relaxation of the cervix with dilation of the cervical os.
- Approaching ovulation, the cervix becomes very relaxed and its protrusion can be seen on the vaginal floor.
- Cervical folds are oedematous
- Vaginal folds glisten with clear lubricant mucus.
Bitch
- Long period of proestrus where the bitch is attractive, but will not allow mating.
- Characterized by:
- Vulval swelling
- Presence of serosanguinous discharge
- Characterized by:
- Followed by standing oestrus:
- Standing to be mated
- Deviation of the tail
- Discharge becomes paler in colour
- Reduction in vulval oedema
- Poor relationship between time of ovulation and external appearance.
Clinical Methods and Assessments for Oestrus Detection
- Assessment of vulval softening
- Assessment of vaginal epithelial cells
- Examination of the appearance of the vaginal wall.
- Measurement of plasma progesterone concentration (detecting a rise in progesterone that preceeds ovulation in this species).
Queen
- Oestrus detection is important, for although the queen is an induced ovulator (thus ovulation will not be missed), the LH response to mating decreases with each day of oestrus and follicles ultimately regress.
- Proestrus is short and often unnoticed.
- Oestrus is characterized by increased vocalization,rubbing and rolling.
- Queen is more active and will seek the tom.
- Stands to be mated by lowering her front quarters and extending her hind legs.
- Demonstrates lordosis
- Tail erect and held to one side.
- Slight mucoid vulval discharge.
Oestrus Behaviour - Anatomy & Physiology Learning Resources | |
---|---|
Flashcards Test your knowledge using flashcard type questions |
Equine Reproduction and Stud Medicine Q&A 15 Equine Reproduction and Stud Medicine Q&A 16 |