Twinning in cattle: I. Foundation animals and genetic and environmental effects on twinning rate

Foundation cows were selected using prior records from one of two sources, private herds or other projects at the Research Center. Comparing twinning rates before and after selection, the repeatability was lower for those from the first (.08) than for those from the second (.16) source with a combined value of .12. Realized heritability of single-parity twinning rate estimated from selection of parents and response in daughters of foundation females was .06. Paternal half-sib estimate of heritability of twinning rate was .02 +/- .07. Estimates of repeatability computed from calving records of females born in the project indicate that permanent environmental effects on twinning rate in cattle are small. Mean calving rate of females born in the project was 1.11 in the data set that included all data and 1.09 in the data set that excluded females from highly selected parents. Twinning rate was greater (P less than .05) in fall than in spring calving (1.13 vs 1.06). Data on twinning rate of a sample of the Swedish Friesian breed were summarized. Mean twinning rate of the Swedish Friesian breed is 2.57%. Age adjusted mean twinning rate of daughters of 32 half-sib sons of one particular Swedish Frieisan sire averaged 5.4% and ranged from .9% to 13.6%. There was no indication of a bimodal distribution, which would be expected if a single gene with a major effect on twinning rate were segregating. The estimated genetic standard deviation (sigma-xn) for mean twinning rates of the 32 sire progeny groups was 1.8%. Observed range among son progenies was .127 or 7.2 sigma-xn, in reasonable agreement with the hypothesis that twinning rate in this population is inherited as a quantitative trait.     

Embryo transfer and twinning in cattle

Recent developments in cattle embryo transfer are discussed, particularly those that are relevant to the commercial application of the technique. Cattle twinning for beef production could offer impressive economic advantages where nutrition is not a limiting factor and intensive management is possible. Evidence is discussed which suggests that cattle twins induced by embryo transfer may hold several advantages over naturally-occurring twins.

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Kurzfassung Neuere Entwicklungen auf dem Gebiet des Embryotransfer beim Rind werden diskutiert. Besonderer Wert wird dabei auf Techniken gelegt, die sich für eine kommerzielle Anwendung eignen. Zwillingsgeburten beim Rind für die Fleischproduktion bietet beeindruckende ökonomische Vorteile in Beständen, wo Ernährung kein limitierender Faktor und intensives Management möglich ist. Es werden Daten diskutiert, die darauf hinweisen, daß Zwillings-kälber von Embryotransferstudien eine Reihe von Vorteilen gegenüber natürlich geborenen Zwillingen aufweisen.

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Resume Les dévelopments récents en matière de transplantation embryonnaire chez les bovins sont discutés en particulier ceux ayant une appliavantages économiques là où la nutrition n'est pas un facteur limitant et l'élevage intensif possible. Il est clairement montré que les jumeaux obtenus par transplantation ovulaire offrent plus d'intérêt que les jumeaux spontanés.

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Riassunto Vengono discussi i recenti sviluppi delle tecniche di transferimento degli embrioni nel bovino, soprattutto di quelli che sono importanti ai fini dell'applicazione commerciale. La produzione di gemelli per una più alta produzione carnea potrebbe offrire rilevanti vantaggi economici dove l'alimentazione non rappresenta un fattore limitante e dove è possible l'allevamento intensivo. Vengono anche discusse le prove secondo cui i parti gemellari indotti nel bovino mediante transferimento embrionale potrebbero presentare molti vantaggi rispetto ai parti gemellari naturali.

     

Association of a single nucleotide polymorphism in SPP1 with growth traits and twinning in a cattle population selected for twinning rate

Continued validation of genetic markers for economically important traits is crucial to establishing marker-assisted selection as a tool in the cattle industry. The objective of the current study was to evaluate the association of a SNP (T(9)/T(10)) in the osteopontin gene (SPP1) with growth rate in a large cattle population spanning multiple generations and representing alleles from 12 founding breeds. This population has been maintained at the US Meat Animal Research Center since 1981 and subjected to selection for twinning rate. Phenotypic records for this population included twinning rate and ovulation rate, providing an opportunity to examine the potential effects of SPP1 genotype on reproductive traits. A set of 2,701 animals was geno-typed for the T(9)/T(10) polymorphism at SPP1. The geno-typic data, including previously genotyped markers on chromosome 6 (BTA6), were used in conjunction with pedigree information to estimate genotypic probabilities for all 14,714 animals with phenotypic records. The genotypic probabilities for females were used to calculate independent variables for regressions of additive, dominance, and imprinting effects. Genotypic regressions were fit as fixed effects in a mixed model analysis, in which each trait was analyzed in a 2-trait model where single births were treated as a separate trait from twin births. The association of the SPP1 marker with birth weight (P < 0.006), weaning weight (P < 0.007), and yearling weight (P < 0.003) was consistent with the previously reported effects of SPP1 genotype on yearling weight. Our data supports the conclusion that the SNP successfully tracks functional alleles affecting growth in cattle. The previously undetected effect of the SNP on birth and weaning weight suggests this particular SPP1 marker may explain a portion of the phenotypic variance explained by QTL for birth and HCW on BTA6.     

Initial results of genomic scans for ovulation rate in a cattle population selected for increased twinning rate

Genomic scans were conducted with 273 markers on 181 sires from a cattle population selected for increased twinning rate to identify chromosomal regions containing genes that influence ovulation rate. Criteria used for selecting markers were number of alleles, ease of scoring, and relative position within linkage group. Markers were multiplexed or multiple-loaded on the gels to reduce the costs and labor required to obtain genotypic data. This approach reduced the number of gels by 45% when compared with running each marker independently. Male animals selected for the genomic scan sired the majority of the population. A modified interval analysis was used in a granddaughter design to compare effects of each allele within sire for 10 different sire families. The midparent deviation of the son's estimated breeding value for ovulation rate was used as the phenotype. Forty-one potential peaks were identified with a nominal significance level < or = 0.05. The 10 peaks with the highest significance levels (P < 0.02) were selected for further analysis. Markers were genotyped across daughters of the sire where nominal significance was found for each of the 10 peaks. One peak (BTA5, relative position 40 cM) was found to be nominally significant in the daughters. The nominal significance levels were P = 0.01 for the sons (n = 32) and P = 0.02 for the daughters (n = 94) of sire 784403. A combined genomewide significance value (P = 0.07) was calculated that accounted for the 10 analyses with sons and the 10 analyses with daughters. These results strongly suggest that this region contains a gene(s) that is involved in the follicular recruitment and development process.     

Ovulation rate and egg recovery in cattle treated repeatedly with pregnant mare serum gonadotrophin and prostaglandin.

Fourteen heifers were superovulated five to 10 times using pregnant mare serum gonadotrophin (PMSG) and prostaglandin in a standardised regime which resulted in a mean interval of 42.1 days (SEM +/- 0.97) between treatments. There was a significant (P less than 0.02) fall in the mean ovulation rate at the second treatment when compared with the first but no significant difference between mean ovulation rate at the second and subsequent treatments. No significant difference between treatments in mean egg recovery rate or in the percentage of normal eggs recovered was demonstrated. Four heifers treated 10 times continued to respond with multiple ovulations on most occasions. At the end of the experiment 13 of the 14 heifers were inseminated and 12 conceived, requiring a mean of 1.67 services per pregnancy.     

Estimates of heritabilities and correlations of traits associated with pelvic area in beef cattle

Pelvic measurements, cow weights and cow ages were obtained on 703 Angus- and Hereford-sired cows from five Louisiana Agricultural Experiment Station herds. Cows were either purebred or crossbred, ranging in age from 1 to 14 yr, and sired by 52 Angus and 63 Hereford bulls. All pelvic measurements were obtained via the rectum by the same technician. Paternal half-sib heritability estimates and genetic (rG) and phenotypic (rP) correlations were computed for pelvic height (PH), pelvic width (PW), pelvic area (PA-I; the product of PH X PW), the ratio of PH to PW and cow weight (CW). Pelvic area was also calculated as an ellipse using the formula PA-II = pi (PH/2)(PW/2). Mean PA-I was 298.5 cm2 while PA-II averaged 234.4 cm2. The pooled heritability estimate for PA-I was .68 +/- .34, indicating that pelvic area is a highly heritable trait and should respond to selection. The estimate for PA-II was similar (.66 +/- .34). The heritability of PW was higher than for PH or PA-I. The heritability of CW was .57 +/- .34 and CW was positively correlated (rG2 = .47 and rP = .40) with PA-I. Direct selection for PA-I was estimated to yield a response of 12.2 cm2 in one generation with a correlated response for CW of 12.5 kg. If change in CW was held at zero using a restricted selection index, about 90% as much increase in PA-I was estimated compared with ignoring CW in the index. Therefore, selection for increased pelvic area can be accomplished without causing large increases in cow size. This should aid in reducing calving difficulty.     

Ovulation and embryonic developmental rate following hCG-stimulation in sows

The hCG induced ovulation in sows was studied by use of ultrasonography, and an investigation of the development and diversity of the zygotes/embryos was performed at 24 h after ovulation. Crossbred sows (N = 48) were weaned (day 0) and checked for heat twice daily from day 3 onwards. From day 4, the ovaries were transrectally scanned twice daily. On day 4, the sows were given an injection of 750 iu hCG i.m. and inseminated 27 +/- 2 h (X +/- SD) and 38 +/- 1 h later. From 38 to 48 h after the hCG injection, the ovaries were scanned at 60 to 90 min intervals. At 24 h after ovulation the oviducts were surgically flushed in 18 sows. Out of the 48 sows, 34 showed heat at 12-36 h after the hCG-treatment and 14 showed heat before the hCG treatment. In the former group of sows, 20 (59%) ovulated within the interval of 38 to 48 h after the hCG treatment, and the follicular size immediately before ovulation was 7.8 +/- 0.6 mm. Among the sows which showed heat before hCG treatment only 7 (50%) ovulated within the above interval and the preovulatory follicle size was larger (8.3 +/- 0.5, p < 0.05) than in the former group of sows, which showed heat after the hCG treatment. The flushing of 18 sows yielded a total of 243 ova, 70 (29%) 1-cell stages, 160 (66%) 2-cell stages and 13 (5%) 4-cell stages. A pronounced difference in the degree of variation in embryonic development was seen between sows: 4 animals yielded 1- to 4-cell stages, one exclusively 2-cell stage. In conclusion, the control of ovulation in sows by hCG treatment will affect the follicular growth and the exact timing of ovulation can not always be relied on. It is strongly recommended to use ultrasonography to monitor the time of ovulation if this parameter is important. Ova recovered at 24 +/- 1 h after the median time of ovulation revealed a pronounced diversity (1- to 4-cell stage) within sows. No obvious relation with this embryonic diversity and the follicular size at ovulation was seen in these data.     

Twinning rate in Norwegian cattle: frequency, (co)variance components, and genetic trends

The overall twinning rate was shown to increase from .6% in the first parity to 4.0% in the sixth parity, and a positive phenotypic trend for twinning rate was observed during the time period considered (1978 to 1995). The distribution of bulls according to the average percentage of multiple births of daughters in first and second parity showed that some bulls had an extremely low twinning frequency and others exceeded the population mean by approximately six times. (Co)variance components were estimated for twinning in first and second parities using a linear sire model. The analysis included either first- and second-crop daughters (1.4 and .9 million records for first- and second-parity cows, respectively) or first-crop daughters only (.6 and .4 million records for first- and second-parity cows, respectively) from 2,043 sires. Heritability estimates were .7 to .8% in the first parity and 2.8% in the second parity. The genetic correlation between twinning in the first and second parities was approximately 1. Part of the phenotypic trend observed can be explained by a genetic trend for twinning rate.     

Use of embryo transfer to induce twinning in beef cattle: embryo survival rate, gestation length, birth weight and weaning weight of calves

Experiments were conducted in 1985 and 1986 at the Eastern Ohio Resource Development Center, Belle Valley, to examine the feasibility of using embryo transfer to induce twinning and to examine the influence of twinning on traits of the cow and calf. Embryos were collected from a total of 14 superovulated Angus donors on two dates each in 1985 and 1986 and were transferred to Angus recipients. A total of 124 embryos were transferred to 79 recipients, with 43 (34.7%) calves born alive. Seven of 45 (15.6%) recipients implanted with two embryos produced twins. In no case did both halves of the 15 embryos that were split to produce identical twins and implanted in the same recipient survive to birth. Proportion of calves born alive did not differ among transfer codes 3 (nonsplit embryos from two different donors implanted in separate uterine horns of the same recipient), 6 (nonsplit embryos from one embryo flush implanted in separate uterine horns of the same recipient) and 7 (nonsplit embryos from two different donors implanted in the same uterine horn of one recipient). Surgical transfers tended to result in a higher proportion of embryos surviving to birth (.43 vs .21; P = .16) and a higher twinning rate (.29 vs .04; P = .36) than did nonsurgical transfers. Age of recipient did not influence embryo survival (P = .98) or twinning rate (P = .99). Gestation length was 5 d shorter (P less than .01) for twin calves than for singles. Singles were 9 kg heavier (P less than .01) at birth and 32 kg heavier (P less than .01) at weaning than twins. However, cows raising twins produced 108 kg (51%) more total weaning weight than did cows raising singles.     

Evaluation of numbers of microscopic and macroscopic follicles in cattle selected for twinning

We hypothesized that the number of microscopic follicles present in the ovaries of cattle selected for twin births (Twinner) would be greater than in the ovaries of contemporary Controls. Ovaries were collected from seven Control and seven Twinner cows at slaughter. The number of Small (1 to 3.9 mm), Medium (4 to 7.9), and Large (> 8 mm) surface follicles was counted and one ovary was fixed for histological evaluation. Fifty to sixty consecutive 6-microm slices were taken from a piece of cortical tissue, approximately 1 cm x 1 cm in area, located between the surface follicles. Microscopic follicles were classified as primordial (oocyte surrounded by a single layer of squamous pregranulosa cells), primary (oocyte surrounded by a single layer of one or more cuboidal granulosa cells), secondary (oocyte surrounded by two or more layers of granulosa cells), or tertiary (oocyte surrounded by multiple layers of granulosa cells with initiation of antrum formation to < or = 1 mm in diameter). The total number of follicles was counted in 200 fields (2 mm x 2 mm) per ovary. A field containing no follicles was classified as empty. There were significantly more secondary follicles in Twinner compared with Control ovaries (12.9 vs 6.3; P < .05). Twinners also tended to have more small surface follicles (35.4 vs 49.0; P < 0.1). We conclude that ovaries of Control and Twinner cows do not differ in the number of primordial follicles or in the number of follicles activated into the growing pool; however, Twinner cows are able to maintain more growing follicles at the secondary and subsequent stages of development.     

Estimation of variance and covariance components to determine heritabilities and repeatability of weaning weight in American Simmental cattle

Components of (co)variance for weaning weight were estimated from field data provided by the American Simmental Association. These components were obtained for the observational components of variance corresponding to a sire, maternal grandsire, and dam within maternal grandsire model. From these estimates, direct additive genetic variance (Sigma2A), maternal additive genetic variance (Sigma2M), covariance between direct and maternal additive genetic effects (SigmaAM), variance of permanent environment(Sigma2pe) and temporary environment variance(Sigma2te) were determined. A procedure to approximate restricted maximum likelihood (REML) estimates of the observational components of variance based on the expectation-maximization (EM) algorithm is described. From these results, phenotypic variance ( ) of weaning weight was 667.88 kg2. Values forSigma2A, Sigma2M, Sigma2pe and Sigma2te were 79,30,58,38,49.45, and 469.97 kg2, respectively. Genetic correlation between direct and maternal additive genetic effects was .16.     

Preimplantation genetic diagnosis in cattle: a review

Preimplantation Genetic Diagnosis (PGD) is reviewed and novel fields where it may be applied are investigated. Technical advances of PGD in cattle embryos have already enabled its integration as a part of the MOET (Multiple Ovulation Embryo Transfer) breeding system. PGD for well-defined selection targets can enhance cattle breeding and embryo trade. It allows embryo selection according to their sex, and it may be used to breed special cow lines, or top bulls, by selecting embryos for valuable production traits using Marker Assisted Selection (MAS). A good allelic profile and/or the insertion of a transgene can be detected by PGD. This review article presents the technical requirements for PGD, and shows that this biotechnological method has great economic potential.     

Moxidectin in cattle: correlation between plasma and target tissues disposition

The time of parasite exposure to active drug concentrations determines the persistence of the antiparasitic activity of endectocide compounds. This study evaluates the disposition kinetics of moxidectin (MXD) in plasma and in different target tissues following its subcutaneous (s.c.) administration to cattle. Eighteen male, 10-month old Holstein calves weighing 120-140 kg were subcutaneously injected in the shoulder area with a commercially available formulation of MXD (Cydectin 1%, American Cyanamid, Wayne, NJ, USA) at 200 micrograms/kg. Two treated calves were killed at each of the following times post-treatment: 1, 4, 8, 18, 28, 38, 48, 58 and 68 days. Abomasal and small intestine mucosal tissue and fluids, bile, faeces, lung, skin and plasma samples were collected, extracted, derivatized and analysed to determine MXD concentrations by high performance liquid chromatography (HPLC) with fluorescence detection. MXD was extensively distributed to all tissues and fluids analysed, being detected (concentrations > 0.1 ng/g; ng/mL) between 1 and 58 days post-treatment. MXD peak concentrations were attained during the first sampling day. MXD maximum concentration (Cmax) values ranged from 52.9 (intestinal mucosa) up to 149 ng/g (faeces). The mean residence time (MRT) in the different tissues and fluids ranged from 6.8 (abomasal mucosa) up to 11.3 (bile) days. MXD concentrations in abomasal and intestinal mucosal tissue were higher than those detected in plasma; however, there was a high correlation between MXD concentrations observed in plasma and those detected in both gastrointestinal mucosal tissues. MXD concentrations were markedly greater in the mucosa than in its respective digestive fluid (P < 0.01). MXD concentrations in skin were higher than those found in plasma (P < 0.01). Drug concentrations recovered in the dermis were greater than those detected in the hypodermal tissue (P < 0.05). Large concentrations of MXD were excreted in bile and faeces. These findings may contribute to an understanding of the relationship between the kinetic behaviour and the persistence of the antiparasite activity of MXD against different ecto-endoparasites in cattle.     

Effects of twinning on postpartum reproductive performance in cattle selected for twin births

The effects of twinning, dystocia, retained placenta, and body weight on postpartum reproduction were evaluated for 3,370 single and 1,014 twin births. Females were bred by AI for 40 d followed by 20 or 30 d of natural service with equal numbers bred and calved in spring and fall. Percentage of dams cyclic by the end of the AI period was lower (P<.05) for dams birthing and nursing a single calf (92.4%) than for dams birthing twins and nursing zero (98.7%) or two (94.7%) calves. Whereas the interval from parturition to first estrus was shorter (P<.01) for dams birthing and nursing a single (56.9 d) than for dams birthing twins and nursing one (68.5 d) or two (69.6 d) calves, length of the interval was further reduced by dystocia in nonlactating dams of either twins or singles (type of birth x dystocia, P<.05). Ensuing pregnancy rates were also affected by type of birth and dystocia. Without dystocia, dams birthing and nursing a single calf had a higher pregnancy rate (79.2%) than dams birthing twins and nursing one (61.7%) or two (66.3%) calves, whereas the lower ensuing pregnancy rates associated with dystocia in dams of singles (71.9%) resulted in similar rates among dams of singles and twins with dystocia (type of birth x dystocia; P<.01). Having a retained placenta resulted in a lower incidence of (93.5 vs. 96.4%, with vs. without; P<.05) and a longer interval to (64.7 vs. 59.2 d; P<.01) estrus while reducing subsequent pregnancy rates (X = 9.6%) in 3 of the 7 yr evaluated (retained placenta x year, P<.01). Because all parous females were bred during the same calendrical period, the shorter gestation length for twin calves (275.6 vs. 281.3 d) resulted in a longer interval from parturition to conception for twin births, whereas means for conception date differed by only 2 d between dams of twins and singles. Furthermore, a reduction (P<.01) in the interval to conception occurred with dystocia in dams of singles (89.3 vs. 85.0 d, without vs. with dystocia) and of twins nursed by zero (116.9 vs. 83.5 d), one (100.2 vs. 92.8 d), or two (96.1 vs. 97.2 d) calves. Another detriment to fertility was the higher incidence of fetal mortality or abortions associated with twin vs. single pregnancies (12.4 vs. 3.5%; P<.01). However, despite the lower conception rates for dams of twins, the increased prolificacy provides an opportunity to increase total beef production with a twinning technology.     

Scrotal circumference in yearling Hereford bulls: adjustment factors, heritabilities and genetic, environmental and phenotypic relationships with growth traits

Field data on 4,233 yearling Hereford bulls were analyzed using fixed and mixed model least-squares procedures to examine factors affecting scrotal circumference; determine appropriate adjustment factors; and study genetic, environmental and phenotypic relationships among scrotal circumference and growth traits. Scrotal circumference was affected by postweaning feed level; contemporary group/feed level; age of dam; and covariates age, weight and height. Of the three covariates, weight had the greatest effect, and any factor which caused an increase in weight tended to increase scrotal circumference. Quadratic effects of age, weight, height and age X age of dam interaction effects were significant or approached significance, but were of minor importance. Large contemporary group effects suggested the need for expressing scrotal circumferences as trait ratios or as deviations from contemporary group means. Scrotal circumference adjustment factors recommended for yearling Hereford bulls were .026 cm X d-1 of age and .8, .2 and .1 for sons of 2-, 3- and 4-yr old dams, respectively. Heritability of weight-adjusted scrotal circumference was .46 +/- .06 compared with .49 +/- .06 for age-adjusted scrotal circumference, indicating considerable additive genetic variation for relative scrotal size. Correlations between scrotal circumference and growth traits were moderate to high. The genetic correlation between scrotal circumference and yearling weight was the highest of these at .44 +/- .16. Potential implications of this relationship are discussed.     

Dwarfism in Hereford cattle: a genetic morphological and biochemical study

A study of 40 Hereford cattle dwarfs in New Zealand confirmed that dwarfism in this country was morphologically the same as that described in North American Herefords and that its mode of inheritance was as an autosomal recessive trait. The histological architecture of growth plates was essentially normal but palisading columns were shorter and more irregular than in controls. A small proportion of chondrocytes in dwarf cartilage showed increased areas of cytoplasmic metachromasia, which probably coincided with increased cystic dilations of endoplasmic reticulum containing granular material as noted by electronmicroscopy. The above growth-plate abnormalities were neither prominent nor consistent enough to be of diagnostic significance; nor do they currently help understanding of the underlying pathogenic mechanism of dwarfing. Histological and mucopolysaccharide excretion studies unequivocally demonstrated that this disease is not a mucopolysaccharidosis as had previously been reported.     

Ovulation rate and litter size in gilts immunized against androstenedione and 17alpha-hydroxyprogesterone

Two experiments were conducted to evaluate the effects of the immunization of gilts against ovarian steroids on ovulation rate and litter size. In Exp. 1, gilts (n = five gilts/treatment) at 165+/-1.6 d of age were immunized against either carrier (Control), androstenedione, or 17alpha-hydroxyprogesterone. Age at puberty and estrous cycle length averaged 208+/-5.5 (P = 0.67) and 20.3+/-2.8 d (P = 0.41), respectively, and were not affected by treatment. The androstenedione- and 17alpha-hydroxyprogesterone immunized gilts had higher (P < 0.02) ovulation rates than Controls (14.2, 14.2, and 11.4+/-0.8, respectively). Total pigs born (P = 0.66) and pigs born live (P = 0.65) for the androstenedione-treated group were not different from Controls. Gestation length was not different (P = 0.36) between any of the treatments and the Controls (115+/-0.9 d). Procedures used in Exp. 2 were similar to those in Exp. 1, except that only Control (n= 18) and 17alpha-hydroxyprogesterone (n = 16) treatments were included and only litter size at farrowing was measured. Total pigs and pigs born live were higher in the 17a-hydroxyprogesterone-treated gilts than in the Controls (12.6 vs 10.5+/-0.6, P < 0.02; and 11.4 vs 9.2+/-0.6; P < 0.01, respectively). Data from this study indicate that litter size in gilts can be increased by immunization against 17alpha-hydroxyprogesterone.     

Dwarfism in Hereford cattle: a genetic morphological and biochemical study

A study of 40 Hereford cattle dwarfs in New Zealand confirmed that dwarfism in this country was morphologically the same as that described in North American Herefords and that its mode of inheritance was as an autosomal recessive trait. The histological architecture of growth plates was essentially normal but palisading columns were shorter and more irregular than in controls. A small proportion of cbondrocytes in dwarf cartilage showed increased areas of cytoplasmic metachromasia, which probably coincided with increased cystic dilations of endoplasmic reticulum containing granular material as noted by electronmicroscopy. The above growth-plate abnormalities were neither prominent nor consistent enough to be of diagnostic significance; nor do they currently help understanding of the underlying pathogenic mechanism of dwarfing. Histological and mucopolysaccharide excretion studies unequivocally demonstrated that this disease is not a mucopolysaccharidosis as had previously been reported.