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ADDENDUM I 228 Slow cooling injury can be caused by solute toxicity (mechanisms not yet elucidated) or by shrinkage of the cells resulting from the hypertonic extracellular solution. Intracellular ice formation (rapid cooling injury) occurs when a cell is not able to maintain osmotic equilibrium with the external environment (Fig. 2). Due to the sudden exaggerated extracellular ice formation, the increase in solutes is so dramatically that the cell cannot respond to it with exosmosis. The cytoplasm will consequently become increasingly supercooled, as such increasing the likelihood of intracellular ice formation (Muldrew et al., 2004). There are multiple hypotheses that attempt to explain how extracellular ice interacts with the plasma membrane in the initiation of intracellular ice formation, but they are all topic of debate. The nature of cellular injury caused by freezing and thawing is very complex. Amann and Pickett (1987) assumed that for stallion spermatozoa, in an extender of given composition, there should be a cooling rate that maximizes sperm survival. Figure 3 represents such a theoretical cooling curve, depicting the sperm survival as a function of the cooling rate. Slow cooling (pathway A to B) will cause sperm damage due to excessive dehydration (solution effect), while rapid cooling (pathway C to D) will cause sperm damage because of intracellular ice formation. The optimal cooling rate is between B and C, and so far this optimal cooling rate can only be determined empirically (Amann and Pickett, 1987). A theoretical approach for determining the optimal cooling rate for the cryopreservation of bull semen has been proposed based on the different compositions in the extender used (Woelders and Chaveiro, 2004). So far, no such work has been presented for stallion spermatozoa. A very remarkable observation concerning cryo-injury was the demonstration that membrane integrity of ram sperm during cryopreservation clearly retained throughout the freeze- thaw procedure, and that the permeabilization occurred not only after the samples had been thawed, but once they reached threshold temperatures (Holt et al., 2005). These findings are in contrast with the concept that the lethality is situated in the intermediate zone of temperature (between freezing point and -60°C) that the cells must traverse twice during the freeze and thaw cycle (Mazur, 1963).
ADDENDUM I Fig. 3. Theoretical curve indicating the survival of cells as function of the cooling rate at the presence (green line) or absence (red line) of glycerol in the extender (Adapted from Amann and Pickett, 1987). All the above clearly states that cryopreservation (of sperm) induces many important changes with the cells. So far the knowledge concerning these changes is incomplete, and therefore till date, semen cryopreservation is a science as well as an art. References Amann R.P., Pickett B.W. 1987. Principles of cryopreservation and a review of cryopreservation of stallion spermatozoa. Journal of Equine Veterinary Science 7:145-173. Hammerstedt R.H., Graham J.K., Nolan J.P. 1990. Cryopreservation of mammalian sperm: What we ask them to survive. Journal of Andrology 11:73-88. Holt W.V., Medrano A., Thurston L.M., Watson P.F. 2005. The significance of cooling rates and animal variability for boar sperm cryopreservation: insights from the cryomicoscope. Theriogenology 63:370-382. Mazur P. 1963. Kinetics of water loss from cells at subzero temperatures and the likelihood of intracellular freezing. Journal of General Physiology 47:347-369. Mazur P. 1984. Freezing of living cells: Mechanisms and implications. American Journal of Physiology 247:125-142. Mazur P., Leibo S.P., Chu E.H. 1972. A two-factor hypothesis of freezing injury. Evidence from Chinese hamster tissue-culture cells. Experimental Cell Research 71:345-355. 229
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AUTOMATED AND STANDARDIZED ANALYSIS
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AUTOMATED AND STANDARDIZED ANALYSIS
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A Area AI Artificial Insemination A
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PREFACE The first recorded case of
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GENERAL INTRODUCTION CHAPTER 1 3
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1.1.1. Introduction CHAPTER 1.1 The
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1.1.4. Concentration CHAPTER 1.1 Ac
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Fluorescent Activated Cell Sorter C
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CHAPTER 1.1 preparation (D, µm) is
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CHAPTER 1.1 These chambers are freq
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CHAPTER 1.1 it is not possible to o
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CHAPTER 1.1 overview of common abno
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CHAPTER 1.1 head, (B5) pyriform hea
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CHAPTER 1.1 The acrosome can be eva
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CHAPTER 1.1 The sperm chromatin str
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Collection and processing of equine
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CHAPTER 1.2 30 (a) Colorado State U
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CHAPTER 1.2 can be induced pharmaco
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CHAPTER 1.2 centrifugation time and
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CHAPTER 1.2 36 → Colloid centrifu
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CHAPTER 1.2 of a stallion stored fo
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CHAPTER 1.2 1.2.4. Cryopreservation
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CHAPTER 1.2 0.5 mL straws, and glyc
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CHAPTER 1.2 determined if the final
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CHAPTER 1.2 46 Chemically defined e
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CHAPTER 1.2 48 Cryopreservation - t
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1.3.1. Standards for equine AI dose
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CHAPTER 1.3 The use of density grad
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CHAPTER 1 Avery B., Greve T. 1995.
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CHAPTER 1 Cheng F.-P., Fazeli A., V
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CHAPTER 1 Flipse R.J., Patton S., A
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CHAPTER 1 Kuisma P., Andersson M.,
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CHAPTER 1 Medeiros A.S.L., Gomes G.
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CHAPTER 1 Pickett B.W. 1993. Collec
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CHAPTER 1 Taberner E., Morató R.,
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CHAPTER 1 Waite J.A., Love C.C., Br
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CHAPTER 2 As evidenced in the gener
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3.1 Influence of technical settings
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CHAPTER 3.1 a CEROS (Hamilton-Thorn
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CHAPTER 3.1 Progressive Motility CA
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CHAPTER 3.1 than at least they shou
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3.2.1. Abstract CHAPTER 3.2 Sperm m
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CHAPTER 3.2 depth but not the chamb
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3.2.3.3. The different chambers and
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CHAPTER 3.2 Finally, Pearson coeffi
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90 80 70 60 50 40 30 20 10 0 drop f
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ISAS20rM Makler10r WHO 120 120 120
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60 PM Rapid 50 40 30 20 10 0 CHAPTE
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Table 5. Averages (± standard erro
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CHAPTER 3.2 sperm (tail) which coul
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CHAPTER 3.2 Hoogewijs M.K., Govaere
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CHAPTER 3.2 Spizziri B.E., Fox M.H.
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3.3.1. Abstract CHAPTER 3.3 Routine
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CHAPTER 3.3 morphological abnormali
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CHAPTER 3.3 The extended semen was
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CHAPTER 3.3 Table 1: Average values
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Repeatability Agreement SQA-Ve PM (
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References CHAPTER 3.3 Arunvipas P.
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3.4.1. Abstract CHAPTER 3.4 In this
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CHAPTER 3.4 therefore in these expe
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CHAPTER 3.4 The averages obtained a
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3.4.4.2. Experiment 2 CHAPTER 3.4 F
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References CHAPTER 3.4 Arunvipas P.
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4.1 Influence of different centrifu
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CHAPTER 4.1 4.1.2. Introduction 140
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CHAPTER 4.1 142 4.1.3.3. Semen proc
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CHAPTER 4.1 144 In the experiment 3
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CHAPTER 4.1 centrifugation (T1) and
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CHAPTER 4.1 (non CP vs CP) was sign
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35 B 80 A 33 beat cross frequency (
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CHAPTER 4.1 152 4.1.4.2. EXPERIMENT
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CHAPTER 4.1 Love and coworkers (200
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CHAPTER 4.1 Love C.C., Brinsko S.P.
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4.2.1. Abstract CHAPTER 4.2 The inc
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4.2.3. Materials and methods 4.2.3.
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CHAPTER 4.2 For the SLC group, 15 m
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CHAPTER 4.2 adding 600 µL of AO st
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4.2.3. Results 4.2.3.1. Sperm quali
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CHAPTER 4.2 The aforementioned “T
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CHAPTER 4.2 Sperm yields markedly d
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References CHAPTER 4.2 Barth A.D.,
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CHAPTER 4.2 Waite J.A., Love C.C.,
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Page 203 and 204: Final reflections 5.4 Actual change
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Page 210 and 211: SUMMARY technique that enables proc
Page 212 and 213: SUMMARY from stallions previously c
Page 214 and 215: SAMENVATTING beweeglijkheid (Hoofds
Page 216 and 217: SAMENVATTING de spermacellen gesele
Page 219 and 220: DANKWOORD Het obligatoire dankwoord
Page 221: DANKWOORD dankzij jou en ze zijn er
Page 225: CURRICULUM VITAE Maarten Hoogewijs
Page 228 and 229: BIBLIOGRAPHY Govaere J., Ducatelle
Page 230 and 231: BIBLIOGRAPHY ABSTRACTS AND PROCEEDI
Page 233 and 234: FUNDAMENTAL ASPECTS OF CRYOPRESERVA
Page 235: ADDENDUM I presence of these CPAs a
Page 239 and 240: ADDENDUM II DETAILED LOADING TECHNI
Page 241: In m’n hoofd is alles heel eenvou
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