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Editorial

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4 pages, 217 KiB

Open AccessEditorial

Advances in Dairy Cattle Reproduction—A Foreword

by Fernando López-Gatius

Animals 2024, 14(18), 2650; https://doi.org/10.3390/ani14182650 - 12 Sep 2024

Viewed by 1400

Abstract

Dairy cattle have played an important role in economic development since the beginning of agriculture [...] Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

Research

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13 pages, 1828 KiB

Open AccessArticle

Brilliant Cresyl Blue Negative Oocytes Show a Reduced Competence for Embryo Development after In Vitro Fertilisation with Sperm Exposed to Oxidative Stress

by Lilli Bittner-Schwerda, Carolina Herrera, Sarah Wyck, Eleni Malama, Christine Wrenzycki and Heinrich Bollwein

Animals 2023, 13(16), 2621; https://doi.org/10.3390/ani13162621 - 14 Aug 2023

Cited by 2 | Viewed by 1242

Abstract

The extent of oxidative damage transferred by the damaged sperm to the progeny is likely to be limited by the oocyte’s repair and antioxidative capacity. We aimed to assess the association between Brilliant Cresyl Blue (BCB) staining in oocytes and their competence for [...] Read more.

The extent of oxidative damage transferred by the damaged sperm to the progeny is likely to be limited by the oocyte’s repair and antioxidative capacity. We aimed to assess the association between Brilliant Cresyl Blue (BCB) staining in oocytes and their competence for embryo development after in vitro fertilisation (IVF) with damaged sperm. For this purpose, bovine sperm were incubated without (non-oxidised sperm, NOX S) or with 100 µM H₂O₂ (oxidised sperm, OX S) and were used to fertilise in-vitro-matured bovine oocytes (BCB-pos./BCB-neg.). Unstained oocytes served as controls (US). Development was assessed at 30, 46, 60 h and on Days (D) 7 and 8 after IVF. Total cell number and apoptotic index were analysed in D7 blastocysts. BCB-neg. oocytes showed lower cleavage rates and blastocyst rates than unstained oocytes after IVF with NOX S (p < 0.05). They showed the highest reduction in D7 blastocyst rate upon fertilisation with OX S and showed a delayed embryo development at 46 and 60 h after IVF compared to embryos produced with NOX S (p < 0.05). Total cell number in blastocysts produced with BCB-neg. oocytes was lower (p < 0.05) in the embryos produced with OX S than in embryos after IVF with NOX S. In conclusion, BCB-neg. oocytes have a lower competence to support embryo development after in vitro fertilisation with oxidised sperm. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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12 pages, 1287 KiB

Open AccessArticle

Expression of Locally Produced Adipokines and Their Receptors during Different Physiological and Reproductive Stages in the Bovine Corpus Luteum

by Granit Thaqi, Bajram Berisha and Michael W. Pfaffl

Animals 2023, 13(11), 1782; https://doi.org/10.3390/ani13111782 - 27 May 2023

Cited by 6 | Viewed by 2655

Abstract

This study aimed to determine the gene expression of different local novel adipokines, such as vaspin, adiponectin, visfatin, and resistin, and their known receptors, namely, heat shock 70 protein 5, adiponectin receptor 1, and adiponectin receptor 2, in the bovine corpus luteum (CL) [...] Read more.

This study aimed to determine the gene expression of different local novel adipokines, such as vaspin, adiponectin, visfatin, and resistin, and their known receptors, namely, heat shock 70 protein 5, adiponectin receptor 1, and adiponectin receptor 2, in the bovine corpus luteum (CL) during different phases of the estrous cycle (on days 1–2, 3–4, 5–7, 8–12, 13–18, >18) and pregnancy (at months 1–2, 3–4, 5–7, >7). The mRNA expression was measured by reverse transcription polymerase chain reaction (RT-qPCR). The mRNA expression levels were normalized to the geometric mean of all three constantly expressed reference genes (cyclophilin A, ubiquitin, ubiquitin C). Our findings suggest that adipokines are expressed and present in all investigated groups, and are specifically up- or downregulated during the estrus cycle and during pregnancy. Vaspin and adiponectin levels were upregulated in the middle and late cycle stages. Resistin was abundant during the CL regression stage and in the first months of pregnancy. The specific expression of adipokine receptors indicates their involvement in the local mechanisms that regulate CL function. Further investigations are required to elucidate the regulative mechanisms underlying the different local effects of adipokines on the ovarian physiology of cows. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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11 pages, 1377 KiB

Open AccessArticle

Acid–Base, Gas, Ions, and Glucose Analysis in Follicular Fluid in Holstein-Friesian Dairy Cows Is Associated with the Follicle Size in Poland

by Bartosz Pawliński, Monika Petrajtis-Gołobów, Michał Trela and Olga Witkowska-Piłaszewicz

Animals 2023, 13(10), 1636; https://doi.org/10.3390/ani13101636 - 14 May 2023

Cited by 3 | Viewed by 1508

Abstract

The lack of fertilization and early pregnancy loss is seen in the quality and maturation of the oocytes. The environment of the first divisions and maturation of the oogonia, which is also a reflection of the quality of the oocyte, is the follicular [...] Read more.

The lack of fertilization and early pregnancy loss is seen in the quality and maturation of the oocytes. The environment of the first divisions and maturation of the oogonia, which is also a reflection of the quality of the oocyte, is the follicular fluid (FF). Thus, the purpose of this study was to investigate the variations in parameters such as pH, pCO₂, pO₂, standard HCO₃⁻, actual HCO₃⁻, base excess (BE), base excess of extracellular fluid (BE ecf), ctCO₂, ions: Na⁺, K⁺, actual ionized calcium Ca²⁺, adjusted ionized calcium (at pH 7.4) Ca²⁺ (7.4), Cl⁻, anion gap (AnGap), and glucose in FF collected from different size follicles in dairy cattle. The most apparent differences were related to pH, K⁺, and Ca²⁺ 7.4 concentration in comparison to changes in follicle size (p < 0.05). Several trends were also evaluated as an increase in follicular size was followed by an increase in pH, BE, and Ca²⁺ 7.4 value and a decrease in the K⁺ concentration (p < 0.05). In conclusion, there are changes in FF formularies associated with the size of follicles. However, more research is necessary to establish the reference value, which then could be one of the factors describing the quality of the follicle and the developmental potential of the associated oocyte. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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16 pages, 1256 KiB

Open AccessArticle

Genomic Prediction for Abortion in Lactating Holstein Dairy Cows

by Robert Wijma, Daniel J. Weigel, Natascha Vukasinovic, Dianelys Gonzalez-Peña, Shaileen P. McGovern, Brenda C. Fessenden, Anthony K. McNeel and Fernando A. Di Croce

Animals 2022, 12(16), 2079; https://doi.org/10.3390/ani12162079 - 15 Aug 2022

Cited by 6 | Viewed by 2993

Abstract

Abortion in dairy cattle causes great economic losses due to reduced animal health, increase in culling rates, reduction in calf production, and milk yield, among others. Although the etiology of abortions can be of various origins, previous research has shown a genetic component. [...] Read more.

Abortion in dairy cattle causes great economic losses due to reduced animal health, increase in culling rates, reduction in calf production, and milk yield, among others. Although the etiology of abortions can be of various origins, previous research has shown a genetic component. The objectives of this study were to (1) describe the development of the genomic prediction for cow abortions in lactating Holstein dairy cattle based on producer-recorded data and ssGBLUP methodology and (2) evaluate the efficacy of genomic predictions for cow abortions in commercial herds of US Holstein cows using data from herds that do not contribute phenotypic information to the evaluation. We hypothesized that cows with greater genomic predictions for cow abortions (Z_Abort STA) would have a reduced incidence of abortion. Phenotypic data on abortions, pedigree, and genotypes were collected directly from commercial dairy producers upon obtaining their permission. Abortion was defined as the loss of a confirmed pregnancy after 42 and prior to 260 days of gestation, treated as a binary outcome (0, 1), and analyzed using a threshold model. Data from a different subset of animals were used to test the efficacy of the prediction. The additive genetic variance for the cow abortion trait (Z_Abort) was 0.1235 and heritability was 0.0773. For all animals with genotypes (n = 1,662,251), mean reliability was 42%, and genomic predicted transmitting abilities (gPTAs) ranged from −8.8 to 12.4. Z_Abort had a positive correlation with cow and calf health traits and reproductive traits, and a negative correlation with production traits. Z_Abort effectively identified cows with a greater or lesser risk of abortion (16.6% vs. 11.0% for the worst and best genomics groups, respectively; p < 0.0001). The inclusion of cow abortion genomic predictions in a multi-trait selection index would allow dairy producers and consultants to reduce the incidence of abortion and to select high-producing, healthier, and more profitable cows. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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11 pages, 492 KiB

Open AccessArticle

The Economic Burden of Chromosome Translocations and the Benefits of Enhanced Screening for Cattle Breeding

by Nicole M. Lewis, Carla Canedo-Ribeiro, Claudia C. Rathje, Rebecca L. Jennings, Maxim Danihel, Lisa M. Bosman, Giuseppe Silvestri and Darren K. Griffin

Animals 2022, 12(15), 1982; https://doi.org/10.3390/ani12151982 - 5 Aug 2022

Cited by 5 | Viewed by 2023

Abstract

The cattle breeding industry, through both of its derivatives (dairy and beef), provides 81% of milk and 22% of meat required globally. If a breeding bull is sub-fertile, this impacts herd conception and birth rates, and it is generally accepted that having a [...] Read more.

The cattle breeding industry, through both of its derivatives (dairy and beef), provides 81% of milk and 22% of meat required globally. If a breeding bull is sub-fertile, this impacts herd conception and birth rates, and it is generally accepted that having a proactive genetic screening programme can prevent further losses. Chromosome translocations are the leading genetic cause of infertility in livestock and, in cattle, this extends beyond the classical 1:29 to other Robertsonian translocations (RobTs) and to reciprocal translocations (RECTs). The incidence of both (collectively termed RTs) varies between breeds and herds; however, we estimate that RECTs are, most likely, at least twice as common as RobTs. The purpose of this study was to develop an industry economic model to estimate the financial impact of an RT event at the herd level. If we assume a conservative incidence rate of 0.4% for Rob1:29 with each one impacting the conception rate by 5%, we calculate that actively screening for and removing a Rob1:29 bull could benefit an impacted herd by GBP 2.3 million (approx. USD 2.8 million) over six years. A recently updated screening protocol developed in our lab for all RTs, however (with a projected combined incidence of 1.2%, impacting conception rates by 10%), could benefit an impacted herd by GBP 7.2 million (nearly USD 9 million) for each RT found. For an industry worth USD 827.4 billion (dairy) and USD 467.7 billion (beef), expanding knowledge on incidence and further dissection of the potential costs (financial and environmental) from RTs is essential to prevent further losses. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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9 pages, 252 KiB

Open AccessArticle

Economics of Rebreeding Nonpregnant Dairy Cows Diagnosed by Transrectal Ultrasonography on Day 25 after Artificial Insemination

by Silviu-Ionuț Borş and Alina Borș

Animals 2022, 12(6), 761; https://doi.org/10.3390/ani12060761 - 17 Mar 2022

Cited by 3 | Viewed by 2362

Abstract

Pregnancy rates of Holstein cows showed a substantial decline in the past years, which caused intensive TAI programs for nonpregnant cows to shorten the period between unsuccessful insemination and the next attempt on the same cow. Although many studies examined the improvement in [...] Read more.

Pregnancy rates of Holstein cows showed a substantial decline in the past years, which caused intensive TAI programs for nonpregnant cows to shorten the period between unsuccessful insemination and the next attempt on the same cow. Although many studies examined the improvement in pregnancy rates following TAI, only a few examined the economic impact of such programs. In this study, we look at the feasibility of reproductive programs that included early pregnancy diagnosis performed by transrectal ultrasonography 25 days after artificial insemination (AI) and TAI of nonpregnant cows. This resulted in the following two TAI programs: a modified OvSynch program with a second PGF2α treatment at 24 h interval (GPPG, n = 100) and a modified OvSynch program with an intravaginal progesterone-release device inserted between days 0–7 (PRID + GPPG, n = 100). Cows included in the TAI programs recorded an improvement in the cumulative pregnancy rate (67% vs. 53%; 69% vs. 53%) compared to those in which this strategy was not applied (p < 0.05). An economic analysis was performed using a decision-support tool to estimate the net present value (NPV; USD/cow/year). The analysis revealed a difference in NPV of 89.6 USD/cow/year between the programs (rebreeding the nonpregnant cows following the TAI program vs. AI at detected estrus). In summary, rebreeding the nonpregnant cows after early negative pregnancy diagnosis (25 days after AI) using this strategy can improve the cumulative pregnancy rate and profitability of dairy farms. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

9 pages, 636 KiB

Open AccessArticle

Response to Therapeutic Abortion in Lactating Dairy Cows Carrying Dead Twins during the Late Embryo/Early Fetal Period

by Fernando López-Gatius

Animals 2021, 11(9), 2508; https://doi.org/10.3390/ani11092508 - 26 Aug 2021

Cited by 5 | Viewed by 3000

Abstract

This study examined the dose–response effect of prostaglandin F_2α (PGF_2α) when used to induce abortion in cows with two dead twins at pregnancy diagnosis 28–34 days post-AI (late embryonic period: LE) or at pregnancy confirmation 49–55 days post-AI (early fetal [...] Read more.

This study examined the dose–response effect of prostaglandin F_2α (PGF_2α) when used to induce abortion in cows with two dead twins at pregnancy diagnosis 28–34 days post-AI (late embryonic period: LE) or at pregnancy confirmation 49–55 days post-AI (early fetal period: EF). The study population consisted of 415 cows receiving a single PGF_2α dose (PG1 group: 254 LE and 161 EF cows) and 200 receiving a 1.5 × PGF_2α dose (PG1.5 group: 121 LE and 79 EF cows). The induced abortion rate was significantly lower (chi-square test; p < 0.0001) in the EF (34.6%) than LE cows (88%), and was also reduced (p = 0.001) in the EF PG1 group (28%) compared with the EF PG1.5 group (48.1%). High milk production (≥45 kg) was the only factor influencing the induced abortion rate in LE cows (odds ratio 0.2; p < 0.0001), whereas the odds ratio for induced abortion of PG1.5 cows was 2.3 (p = 0.005) in EF cows. In conclusion, an increased PGF_2α dose had no effect on abortion induction during the late embryonic period and its rate was inversely associated with milk production. This higher PGF_2α dose did, however, offer benefits during the early fetal period. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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Review

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24 pages, 1401 KiB

Open AccessReview

Current Insights in the Repeat Breeder Cow Syndrome

by Carlos Carmelo Pérez-Marín and Luis Angel Quintela

Animals 2023, 13(13), 2187; https://doi.org/10.3390/ani13132187 - 3 Jul 2023

Cited by 12 | Viewed by 5172

Abstract

Cows can have difficulties becoming pregnant, and in certain cases, these reproductive failures do not have an evident cause. Furthermore, when these failures are repeated three or more times with estrous cycles of normal duration and in the absence of evident clinical signs, [...] Read more.

Cows can have difficulties becoming pregnant, and in certain cases, these reproductive failures do not have an evident cause. Furthermore, when these failures are repeated three or more times with estrous cycles of normal duration and in the absence of evident clinical signs, it is considered repeat breeder cow (RBC) syndrome. A substantive incidence of RBC syndrome has been reported all over the world, which severely affects the farm economy. This paper reviews those studies particularly focused on RBC syndrome from 2000 to 2023 but also includes consolidated information until this date. Hormonal imbalances, undetectable oviductal or uterine defects, or poor oocyte or embryo quality have been reported as causes of RBC syndrome, while subclinical endometritis has been considered a relevant causal agent. However, it is unresolved why this condition is recurrent in certain animals, despite the implementation of corrective management actions or treatments. Recent studies evaluate the putative role of certain genes, factors, hormones, or proteins in the pathogenesis of RBC syndrome. Numerous risk factors contribute to the appearance of this syndrome, and some of them could be mitigated to partially prevent this infertility, while others cannot be changed. Due to the complexity of this syndrome, it is important to increase knowledge about the mechanisms involved, develop new diagnostic tools to differentiate causal agents, and implement new treatments to restore fertility. There is consensus about the huge repercussions of this syndrome on farm profitability, but further studies are now needed to describe its economic impact. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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16 pages, 1750 KiB

Open AccessReview

Using Pregnancy-Associated Glycoproteins (PAGs) to Improve Reproductive Management: From Dairy Cows to Other Dairy Livestock

by Olimpia Barbato, Laura Menchetti, Gabriele Brecchia and Vittoria Lucia Barile

Animals 2022, 12(16), 2033; https://doi.org/10.3390/ani12162033 - 10 Aug 2022

Cited by 22 | Viewed by 4069

Abstract

Pregnancy success represents a major issue for the economic income of cattle breeders. Early detection of pregnant and non-pregnant animals, as well as the prediction of early pregnancy failure, can influence farm management decisions. Several diagnostic tools for pregnancy are currently available. Among [...] Read more.

Pregnancy success represents a major issue for the economic income of cattle breeders. Early detection of pregnant and non-pregnant animals, as well as the prediction of early pregnancy failure, can influence farm management decisions. Several diagnostic tools for pregnancy are currently available. Among these, pregnancy-associated glycoproteins (PAGs) have been shown to be useful for identifying the presence of vital embryos and for pregnancy follow-up monitoring. This review presents an overview of the PAGs’ functions, their pregnancy trends, and their use as a tool to improve reproductive management in bovine and other dairy livestock, such as small ruminants and buffalos. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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20 pages, 372 KiB

Open AccessReview

Accuracy to Predict the Onset of Calving in Dairy Farms by Using Different Precision Livestock Farming Devices

by Ottó Szenci

Animals 2022, 12(15), 2006; https://doi.org/10.3390/ani12152006 - 8 Aug 2022

Cited by 9 | Viewed by 2873

Abstract

Besides traditional methods such as evaluation of the external preparatory and behavioral signs, which even presently are widely used also in large dairy farms, there are several new possibilities such as measuring body (intravaginal, ventral tail-base surface, ear surface, or reticulo-ruminal) temperature, detecting [...] Read more.

Besides traditional methods such as evaluation of the external preparatory and behavioral signs, which even presently are widely used also in large dairy farms, there are several new possibilities such as measuring body (intravaginal, ventral tail-base surface, ear surface, or reticulo-ruminal) temperature, detecting behavioral signs (rumination, eating, activity, tail raising) or detecting the expulsion of the device inserted into the vagina or fixed to the skin of the vulva when allantochorion appears in the vulva to predict the onset of the second stage of calving. Presently none of the single sensors or a combination of sensors can predict the onset of calving with acceptable accuracy. At the same time, with the exception of the iVET^® birth monitoring system, not only the imminent onset of calving could be predicted with high accuracy, but a significantly lower prevalence rate of dystocia, stillbirth, retained fetal membranes, uterine diseases/clinical metritis could be reached while calving-to-conception interval was significantly shorter compared with the control groups. These results may confirm the use of these devices in dairy farms by allowing appropriate intervention during calving when needed. In this way, we can reduce the negative effect of dystocia on calves and their dams and improve their welfare. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

9 pages, 1375 KiB

Open AccessReview

Effects of Heat Stress on Follicular Physiology in Dairy Cows

by Fabio De Rensis, Roberta Saleri, Irina Garcia-Ispierto, Rex Scaramuzzi and Fernando López-Gatius

Animals 2021, 11(12), 3406; https://doi.org/10.3390/ani11123406 - 29 Nov 2021

Cited by 20 | Viewed by 3834

Abstract

Follicular organization starts during mid-to-late fetal life with the formation of primordial follicles. The bilateral interplay between the oocyte and adjoining somatic cells during follicular growth and ovulation may be sensitive to heat stress (HS). Mechanisms giving rise to pre-ovulatory temperature gradients across [...] Read more.

Follicular organization starts during mid-to-late fetal life with the formation of primordial follicles. The bilateral interplay between the oocyte and adjoining somatic cells during follicular growth and ovulation may be sensitive to heat stress (HS). Mechanisms giving rise to pre-ovulatory temperature gradients across reproductive tissues are mostly regulated by the pre-ovulatory follicle, and because the cooling of the gonads and genital tract depends on a counter-current transfer system of heat, HS may be considered a major factor impairing ovulation, fertilization and early embryo development. There is evidence of a long-lasting influence of HS on oogenesis and final follicular maturation. Follicular stages that are susceptible to HS have not been precisely determined. Therefore, the aim of this review was to describe the influence of HS during the staged follicular development in dairy cattle, from the activation of primordial follicles to ovulation. Some clinical prospects are also considered. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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Other

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8 pages, 2325 KiB

Open AccessCommentary

Revisiting the Timing of Insemination at Spontaneous Estrus in Dairy Cattle

by Fernando López-Gatius

Animals 2022, 12(24), 3565; https://doi.org/10.3390/ani12243565 - 16 Dec 2022

Cited by 12 | Viewed by 3558

Abstract

Cows show spontaneous estrus over 8–20 h but become refractory to the bull about 10–12 h before ovulation. This indicates that ovulation occurs 10–12 h after the end of estrus behavior, yet spermatozoa from the bull ejaculate need to undergo maturation and capacitation [...] Read more.

Cows show spontaneous estrus over 8–20 h but become refractory to the bull about 10–12 h before ovulation. This indicates that ovulation occurs 10–12 h after the end of estrus behavior, yet spermatozoa from the bull ejaculate need to undergo maturation and capacitation for 6 to 8 h in the female reproductive tract before they are capable of fertilization. Traditionally, the onset of estrus has been considered the best timing for artificial insemination (AI) in cattle, that is, 6 to 24 h from the first signs of estrus. However, recent findings suggest this interval should be reduced to 16 to 6 h before ovulation, bringing it closer to the end of estrus. In this review, the end of estrus rather than its onset is proposed as the best guide for AI timing in dairy cattle, and physiological indicators of late estrus are discussed such as relaxation of the intravaginal part of the uterus, a lower cervical mucus viscosity and a softer pre-ovulatory follicular consistency as simple cues indicating a cow is ready for service. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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5 pages, 243 KiB

Open AccessCommentary

The Uterus as an Influencing Factor for Late Embryo/Early Fetal Loss—A Clinical Update

by Zoltán Szelényi, Levente Kovács, Ottó Szenci and Fernando Lopez-Gatius

Animals 2022, 12(15), 1873; https://doi.org/10.3390/ani12151873 - 22 Jul 2022

Cited by 4 | Viewed by 1983

Abstract

Here we revise circumstances of non-infectious causes in which the uterus may be associated with pregnancy loss during the late embryo/early fetal period (following a positive pregnancy diagnosis in lactating dairy cows). As the uterine size increases with parity and pregnant heifers with [...] Read more.

Here we revise circumstances of non-infectious causes in which the uterus may be associated with pregnancy loss during the late embryo/early fetal period (following a positive pregnancy diagnosis in lactating dairy cows). As the uterine size increases with parity and pregnant heifers with no detrimental effects of a previous parturition, a primigravid uterus is proposed as a reference for identifying risk factors that negatively influence pregnancy in lactating cows. Cows suffering placenta retention or with a large uterus at insemination were selected as topics for this revision. Retained placenta, that occurs around parturition, has a long-lasting influence on subsequent pregnancy loss. Although retained placenta is a particularly predisposing factor for uterine infection, farm conditions along with cow factors of non-infectious cause and their interactions have been identified as main factors favoring this disorder. A large uterus (cervix and uterine horns lying outside the pelvic cavity) with no detectable abnormalities has been associated with low fertility and with a greater incidence of pregnancy loss. A large reproductive tract may well derive from an inadequate uterine involution. Therefore, peripartum management and strategies to reduce the incidence of uterine disorders should reduce their associated financial losses in the herds. Full article

(This article belongs to the Special Issue Advances in Dairy Cattle Reproduction)

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