Growth of the African elephant (Loxodonta africana)

The three-stage desk calculation of the von Bertalanffy equation to describe growth in height and weight with age in the elephant is compared with a new approach to calculating the three coefficients in the function by a computer. The two methods give different results with respect to the weight/age calculations. Theoretical von Bertalanffy equations calculated by both methods to describe growth in height and weight with age in the African elephant in Zambia are compared with previously published equations for the elephant in East Africa. Details are given of growth in height in two known-age African elephants, a female ‘Diksie’ and a male ‘Kartoum’. Theoretical growth in height curves for the female African and Asiatic elephant are compared. The coefficients K and / for growth in height are not transferable to the growth in weight equations. Inherent inaccuracies in the calculation of the coefficients in the von Bertalanffy equation are discussed, and it is concluded that in animals which have a long life-span such as the elephant, the equation serves as a purely empirical representation of weight-at-age data and that there is little biological significance in the parameters it contains. The computer-calculated curves give the best fit to the data. The regression of log age on log shoulder height from 2–20 years of age provides a more realistic approach to comparative growth studies. The increase in adrenal weight with age is linear. Tusk growth in relation to age and sex in Zambia is compared with East Africa. It is concluded that the tusks in Zambia are smaller and are more difficult to sex correctly than their East African counterparts, possibly a consequence of the Zambian elephant having a greater degree of tusk wear. Allometric growth is described with emphasis on the estimation of body weight from shoulder height. The most reliable estimates are obtained from a purely empirical representation of the data, a semilog plot of log body weight on shoulder height.     

Population dynamics of the African elephant (Loxodonta africana)

Recent studies on elephant populations from East Africa and from Zambia have suggested that as population density increases, so does the mean age at puberty and the mean calving interval. At the same time there is also an increase in the proportion of old females that are reproductively inactive. By constructing elephant population models, it is possible to investigate the extent to which these "homeostatic mechanisms" will regulate an elephant population. The models indicate that a change in the duration of the calving interval is more important as a population regulating mechanism than a change in the age at puberty, and that the proportion of old reproductively inactive females is of little significance. The importance of neonatal mortality in controlling population growth is emphasised by the models, and they also show that an annual population increase of 4% would be close to the maximum value.     

Synchronization of defaecation in the African elephant (Loxodonta africana)

The ecological significance of defaecation in African elephants ( Loxodonta africana ) has been discussed from a number of points of view. Dropping counts have been used to investigate population size and movements (Wing & Buss, 1970; Laws, Parker & Johnstone, 1975); the chemical composition of dung has been examined (Dougall, 1963; Weir, 1972); and dung decomposition and its role in nutrient cycles have been described (Anderson & Coe, 1974). Defaecation in captive African elephants has been examined (Coe, 1972) and food consumption of wild elephant herds has been calculated from data on gross assimilation efficiency and dung production (Coe, 1972; Rees, 1982). This paper presents evidence of a synchronization of defaecation in elephants and discusses its possible ecological significance.     

Effect of cryoprotective diluent and method of freeze-thawing on survival and acrosomal integrity of ram spermatozoa

A multifactorial study analyzed the effects of freezing method, cryoprotective diluent, semen to diluent ratio, and thawing velocity on post-thaw motility, progressive status, and acrosomal integrity of ram spermatozoa. Although semen to diluent ratio (1:3 vs 1:6, v/v) had no effect (P greater than 0.05), overall post-thaw spermatozoal viability was highly dependent on freezing method and cryoprotectant. Improved results were obtained by freezing semen in 0.5-ml French straws compared to dry ice pelleting. Manually freezing straws 5 cm above liquid nitrogen (LN2) was comparable to cooling straws in an automated, programmable LN2 unit. Of the two cryoprotective diluents tested, BF5F (containing the surfactant component sodium and triethanolamine lauryl sulfate) yielded approximately 50% fewer (P less than 0.05) spermatozoa with loose acrosomal caps compared to TEST. Thawing straws in a water bath at a higher velocity (60 degrees C for 8 sec) had no effect (P greater than 0.05) on spermatozoal motility, progressive status ratings, or acrosomal integrity when compared to a lower rate (37 degrees C for 20 sec). For the TEST group, thawing pellets in a dry, glass culture tube promoted (P less than 0.05) percentage sperm motility at 3 and 6 hr post-thawing, but for BF5F diluted semen this approach decreased the % of spermatozoa with normal apical ridges. The results suggest that the poor fertility rates often experienced using thawed ram semen likely result not only from reduced sperm motility, but also from compromised ultrastructural integrity. This damage is expressed by an increased loosening of the acrosomal cap, a factor which appears insensitive to freezing method but markedly influenced by the cryoprotective properties of the diluents tested.     

Comparative cytogenetics of the African elephant (Loxodonta africana) and Asiatic elephant (Elephas maximus)

G- and C-banded karyotypes of the two extant species of the mammalian order Proboscidea are presented for the first time. Chromosome complements were 2n = 56 in both Loxodonta africana and Elephas maximus. Comparisons between the species demonstrated a high level of chromosome band homology, with 26 conserved autosomal pairs. The normal diploid karyotype of L. africana had 25 acrocentric/telocentric and two metacentric/submetacentric autosomal pairs. E. maximus differed by having one less acrocentric and one additional submetacentric pair due to either a heterochromatic arm addition or deletion involving autosomal pair 27. Several acrocentric autosomes of L. africana exhibited small short arms that were absent in homologous chromosomes of E. maximus. The X chromosomes in both species were large submetacentric elements and were homologous. However, the small acrocentric Y chromosomes differed; in E. maximus it was slightly larger and had more distinct G-bands than its counterpart in L. africana. Extant Elephantidae appear to be relatively conservative in their rates of chromosomal change compared to some other mammalian families. The high-quality banded karyotypes presented here should prove useful as references in future chromosome analyses of elephant populations and in comparative cytogenetic studies with other ungulate orders.     

Musth and urinary testosterone concentrations in the African elephant (Loxodonta africana)

Urine samples were obtained from free-ranging African elephants that were considered to be in and out of musth. Testosterone concentrations, measured by radioimmunoassay were significantly greater in males that were in or around the time of behavioural musth. This study supports a correlation between the observed behavioural characteristics of musth and urinary testosterone levels.     

Blood characteristics of the African elephant (Loxodonta africana cyclotis).

Mean corpuscular volume, hematocrit, hemoglobin concentration and red and white cell counts from five domesticated African elephants were measured. The results obtained are tabulated and compared with those from shot and chemically immobilized African elephants.     

Structural characterization of oligosaccharides in the milk of an African elephant (Loxodonta africana africana)

The oligosaccharides present in the milk of an African elephant (Loxodonta africana africana), collected 4 days post partum, were separated by size exclusion-, anion exchange- and high-performance liquid chromatography (HPLC) before characterisation by (1)H NMR spectroscopy. Neutral and acidic oligosaccharides were identified. Neutral oligosaccharides characterised were isoglobotriose, Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)Glc, Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)Glc, Gal(alpha1-3)Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)Glc and a novel oligosaccharide that has not been reported in the milk or colostrum of any other mammal: Gal(alpha1-3)Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)Glc. Acidic oligosaccharides that are also found in the milk of Asian elephant were Neu5Ac(alpha2-3)Gal(beta1-4)Glc, Neu5Ac(alpha2-6)Gal(beta1-4)Glc, Neu5Ac(alpha2-3)Gal(beta1-4)[Fuc(alpha1-3)]Glc, Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)Glc, Neu5Ac(alpha2-3)Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)Glc, Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-3)Gal(beta1-4)Glc and Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3){Gal(alpha1-3)Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-6)}Gal(beta1-4)Glc, while Neu5Gc(alpha2-3)Gal(beta1-4)Glc, Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3)Gal(beta1-4)[Fuc(alpha1-3)]Glc, Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3)[Gal(beta1-4)GlcNAc(beta1-6)]Gal(beta1-4)Glc and Neu5Ac(alpha2-6)Gal(beta1-4)GlcNAc(beta1-3){Gal(beta1-4)[Fuc(alpha1-3)]GlcNAc(beta1-6)}Gal(beta1-4)Glc have not been found in Asian elephant milk. The oligosaccharides characterised contained both alpha(2-3)- and alpha(2-6)-linked Neu5Ac residues. They also contain only the type II chain, as found in most non-human, eutherian mammals.     

Circulating oestrogen concentrations during pregnancy in the African elephant (Loxodonta africana)

Oestrone, oestradiol-17 beta and oestriol were measured in plasma samples from non-pregnant and pregnant African elephants shot in the wild. Enzymic hydrolysis of plasma showed that approximately 90 and 96% of the total (i.e. conjugated plus unconjugated) concentrations of oestrone and oestradiol-17 beta, respectively were represented by conjugated hormones. Unconjugated oestrogens remained low (less than 50 pg ml) in all samples, with no distinction between non-pregnant and pregnant animals. Levels of total oestrone during pregnancy varied between 160 and 594 pg/ml but were not significantly different from non-pregnant values. Total oestradiol-17 beta concentrations were significantly elevated during pregnancy (P less than 0 X 01) and, despite considerable individual variation (193-1428 pg/ml), were consistently higher than non-pregnant values after 6 months of gestation. The elevated levels of oestradiol-17 beta resulted in a reversal of the total oestradiol-17 beta: oestrone concentration ratio at about 6 months of pregnancy. Concentrations of total oestriol did not exceed 103 pg/ml. An indirect method of measurement indicated that oestradiol-17 beta sulphate was probably the most abundant circulating oestrogen during pregnancy in the African elephant.     

Lectin histochemistry of the temporal gland of the African elephant (Loxodonta africana)

The study demonstrates the free sugar spectrum of the secretion of the tubuloaveolar temporal gland of the African elephant (Loxodonta africana), using lectin histochemistry. In the elephant, the spectrum contained, besides strongly varying amounts of α-D-mannose, very remarkable reactions for α-D-galactose and to a certain extent also for α-D-N-acetyl-galactosamine or a/?-D-N-acetylglucosamine. This is in contrast to the free sugar spectrum of the secretion of the mammalian apocrine tubular skin glands. Considering also that the production of any water binding mucus seems to be negligible, the variations of the free sugar contents found probably originate from another important task of the secretory cells of the temporal gland. This means that our findings corroborate the view of highly active acquired immunity by an intensive processing and presenting of lipid antigens by dendritic cells (APC), in particular of the CD1 family.     

Milk composition of three free-ranging African elephant (Loxodonta africana africana) cows during mid lactation

Data are presented that indicate the dynamic changes of nutrients in milk from three free ranging African elephant (Loxodonta africana africana) cows during lactation. At the respective collection times of 12, 14 and 18 months of lactation the nutrient content was 47.3, 52.0 and 68.6 g protein; 60.7, 87.4 and 170.8 g fat; 1.6, 2.1 0.5 g lactose and 20.9, 21.5 and 8.6 g oligosaccharides per kg milk. The protein fraction respectively consisted of 18.0, 31.7 and 45.9 g caseins/kg milk and of 29.3, 20.3 and 22.7 g whey proteins/kg milk. Electrophoresis and identification of protein bands showed that polymorphs of one whey protein may be present in elephant's milk similar to polymorphs of alpha-lactalbumin found in cow's milk. From the middle of the lactation time lactose was replaced by oligosaccharides as major carbohydrate, and the major compound of these was identified as isoglobotriose by 1H NMR spectroscopy. The lipid fraction contains a high content, of capric and lauric acids, approximately 70% of the total fatty acids, and low content of myristic, palmitic and oleic acids. During these lactation times the content of short chain fatty acids, capric and caprylic acids increased, while fatty acids lauric acid and longer decreased.     

The temporomandibular joint disc of Asian elephant (Elephas maximus) and African elephant (Loxodonta africana)

The temporomandibular joint (TMJ) is a synovial articulation between the mandibular head of the condylar process of the mandible and the mandibular fossa of the squamous temporal bone. Extensions of the fibrous capsule into the joint space form a biconcave disc that functions as an articulating surface and divides the joint into dorsal and ventral compartments. The TMJ disc plays a crucial role in normal functioning of the joint, and differences in cranial morphology, mastication patterns, and diet are reflected in the material and biochemical properties of the disc. The purpose of the present case study was to compare the regional histologic differences between two elephant genera and quantify the biochemical and biomechanical properties of the African elephant disc. This study provides a unique insight into the elephant TMJ disc and also provides a comparison between the African and the Asian elephant genera. The results demonstrate several remarkable findings. First, structure–function relationships exist within the elephant TMJ disc. Second, regional variations exist in the elephant TMJ disc, and these are likely to correlate with its functional requirement. Additionally, it is apparent that some properties of the disc vary with the specific anatomy, diet requirement, and jaw motion. Finally, in comparison with the TMJ disc of other species, it is clear that, although the elephant disc is unique, it has properties that transcend and are preserved among the species.     

Initial findings on visual acuity thresholds in an African elephant (Loxodonta africana)

There are only a few published examinations of elephant visual acuity. All involved Asian elephants (Elephas maximus) and found visual acuity to be between 8′ and 11′ of arc for a stimulus near the tip of the trunk, equivalent to a 0.50 cm gap, at a distance of about 2 m from the eyes. We predicted that African elephants (Loxodonta africana) would have similarly high visual acuity, necessary to facilitate eye‐trunk coordination for feeding, drinking and social interactions. When tested on a discrimination task using Landolt‐C stimuli, one African elephant cow demonstrated a visual acuity of 48′ of arc. This represents the ability to discriminate a gap as small as 2.75 cm in a stimulus 196 cm from the eye. This single‐subject study provides a preliminary estimate of the visual acuity of African elephants. Zoo Biol 29:30–35, 2010. © 2009 Wiley‐Liss, Inc.     

Circulating progesterone concentrations and ovarian functional anatomy in the African elephant (Loxodonta africana)

Mean plasma progesterone concentrations measured in pregnant and non-pregnant elephants did not differ significantly from each other because of considerable variation, particularly for stage of pregnancy. Maximum progesterone values were recorded during pregnancy (5-8 months) and declined towards term (22 months). The numbers of corpora lutea or total luteal tissue volume were not critical in maintaining progesterone secretion. An increase in plasma progesterone concentrations with the luteal phase of the ovarian cycle was evident. A possible role of the placenta in the second half of gestation is indicated by an increase in fetal progesterone concentrations towards term.     

Milk intake and development in a newborn captive African elephant (Loxodonta africana)

In August 2003 the San Diego Zoo's Wild Animal Park (WAP) and the Lowry Park Zoo, under the auspices of the AZA's Elephant SSP and a USFWS permit, imported 3.8 African elephants (Loxodonta africana) from the Kingdom of Swaziland. When they were captured, transrectal ultrasound examinations revealed that one nulliparous cow was approximately 10 months pregnant. At the time of their arrival (August 2003), all of the animals were estimated to be approximately 13 years old and were thought to be nulliparous. Based on the ultrasound examination results and the average African elephant gestation period, parturition was predicted to be 20 February 2004. In this report, we provide the first detailed data about nursing activity around the clock and newborn calf development, describe maternal and neonatal nighttime activity budgets, and explore maternal weight changes during suckling and lactation. The newborn calf suckled significantly more at night than during the day, but suckled for only about 2 hr per 24‐hr period. Regression analysis revealed that through the first 3 months of life the calf gained 0.385 kg/day while it suckled on a regular basis. We compare our findings with published information on wild elephants, and conclude that although the growth rate is reduced compared to hand‐reared elephant calves, the suckling patterns are almost identical to those reported for wild calves. Zoo Biol 0:1–7, 2005. © 2005 Wiley‐Liss, Inc.     

Histo- and cytophysiology of the lactating mammary gland of the African elephant (Loxodonta africana)

The lactating mammary gland of the African elephant (Loxodonta africana) has been studied with a panel of morphological techniques focusing on (1) the functional changes during the secretory process, (2) proliferative process [by application of proliferating cell nuclear antigen (PCNA) immunohistochemistry] and apoptotic phenomena [by use of the TUNEL technique] in the individual lobules, and (3) components of milk and milk-fat-globule membrane. In the lactating gland, the lobules are variably differentiated; within a lobule, however, the alveoli are usually similarly differentiated. The morphology of their alveoli suggests a classification of the lobules into types 1–3. Lobules of type 1 are composed of immature tubular alveoli with mitotic figures and numerous PCNA-positive nuclei; advanced type 1 alveoli contain abundant glycogen and specific secretory granules. Lobules of type 2 are further subdivided. In type 2a lobules, the epithelial cells of the alveoli form tall apical protrusions, which in part are occupied by small lipid droplets and which are pinched off in an apocrine fashion. The number of lysosomes varies considerably. Type 2b is the most common type, with striking basal membrane foldings, abundant rough endoplasmic reticulum cisterns, large Golgi apparatus, numerous mitochondria, lipid droplets, and protein vesicles with 30- to 90-nm-wide casein micelles. The lipid droplets are pinched off with minimal amounts of cytoplasm. Type 2c is composed of alveoli with a cuboidal epithelium and few signs of secretory activity. Increasing expression of peanut-agglutinin-binding sites parallels the maturation and differentiation of the glandular cells. Type 3 lobules are marked by numerous TUNEL-positive nuclei and large lipid droplets and are apparently degenerating structures. Cytokeratin (CK) 14 is usually present in the myoepithelial cells; CK 19 and CK 7 mark ductal and immature alveolar epithelia. Milk protein content varies between 2.6% and 6.3%, and casein micelles range from 35 to 90 nm in diameter. The diameter of intra-alveolar milk fat globules ranges from 5 to 25 µm and the membranes bear a filamentous surface coat composed of membrane-anchored mucins; gel-electrophoretic analysis of these mucins from different individuals demonstrates the presence of mucin MUC 1, which is expressed with considerable genetic heterogeneity.