Effects of antler breakage on mating behavior in male tule elk (<Emphasis Type="Italic">Cervus elaphus nannodes</Emphasis>)

Although antler size has been identified as a primary determinant of dominance, fighting success, and reproductive success in male cervids, >80% of the male tule elk (Cervus elaphus nannodes) in the Owens Valley, California, experience antler breakage. To determine the effect of antler breakage on male mating success, we recorded antler morphology, body size, and mating behavior of male elk throughout the rut. Antler breakage, regardless of severity, had no effect on male–male assessment, fighting success, or harem-holding status. The factor consistently associated with our indices of male mating success was not antler size but body size. Although antler size is frequently emphasized as a key factor in male dominance and social rank, this association may reflect the correlation between antler size and body size. In the Owens Valley, it appears that male elk are not assessing competitors based on antler morphology but on other characteristics. An erratum to this article can be found at     

Gene expression dynamics in deer antler: mesenchymal differentiation toward chondrogenesis

Annual re-growth of deer antler represents a unique example of complete organ regeneration. Because antler mesenchymal cells retain their embryonic capacity to develop into cartilage or bone, studying antler development provides a natural system to follow gene expression changes during mesenchymal differentiation toward chondrogenic/osteogenic lineage. To identify novel genes involved either in early events of mesenchymal cell specialization or in robust bone development, we have introduced a 3 K heterologous microarray set-up (deer cDNA versus mouse template). Fifteen genes were differentially expressed; genes for housekeeping, regulatory functions (components of different signaling pathways, including FGF, TGFβ, Wnt), and genes encoding members of the Polycomb group were represented. Expression dynamics for genes are visualized by an expression logo. The expression profile of the gene C21orf70 of unknown function is described along with the effects when over-expressed; furthermore the nuclear localization of the cognate protein is shown. In this report, we demonstrate the particular advantage of the velvet antler model in bone research for: (1) identification of mesenchymal and precartilaginous genes and (2) targeting genes upregulated in robust cartilage development. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Identification of key tissue type for antler regeneration through pedicle periosteum deletion

Epimorphic regeneration is the “holy grail” of regenerative medicine. Research aimed at investigating the various models of epimorphic regeneration is essential if a fundamental understanding of the factors underpinning this process are to be established. Deer antlers are the only mammalian appendages that are subject to an annual cycle of epimorphic regeneration. In our previous studies, we have reported that histogenesis of antler regeneration relies on cells resident within the pedicle periosteum (PP). The present study elaborates this finding by means of functional studies involving the deletion of PP. Four yearling and four 2-year-old stags were selected for total PP deletion or partial PP deletion experiments. Of the animals in the total PP deletion group, one showed no signs of antler regeneration throughout the antler growth season. Two showed substantial and one showed marginal delays in antler regeneration (at 34, 20 and 7 days, respectively) compared with the corresponding sham-operated sides. Histological investigation revealed that the delayed antlers were derived from regenerated PP. Unexpectedly, the regenerative capacity of the antler from the total periosteum-deleted pedicles depended on antler length at surgery. Of the four deer that had partial PP deletion, two regenerated antlers exclusively from the left-over PP on the pedicle shafts in the absence of participation from the pedicle bone proper. The combined results from the PP deletion experiments convincingly demonstrate that the cells of the PP are responsible for antler regeneration. The authors thank the New Zealand Foundation of Research, Science and Technology and Deer Industry New Zealand for funding their research.     

Characterization of heparan sulfate from the unossified antler of Cervus elaphus

The antler is the most rapidly growing tissue in the animal kingdom. According to previous reports, antler glycosaminoglycans (GAGs) consist of all kinds GAGs except for heparan sulfate (HS). Chondroitin sulfate is the major antler GAG component comprising 88% of the total uronic acid content. In the current study, we have isolated HS from antler for the first time and characterized it based on both NMR spectroscopy and disaccharide composition analysis. Antler GAGs were isolated by protease treatment and followed by cetylpyridinium chloride precipitation. The sensitivity of antler GAGs to heparin lyase III showed that this sample contained heparan sulfate. After incubation of antler GAGs with chondroitin lyase ABC, the HS-containing fraction was recovered by ethanol precipitation. The composition of HS disaccharides in this fraction was determined by its complete depolymerization with a mixture of heparin lyase I, II, and III and analysis of the resulting disaccharides by the reversed-phase (RP) ion pairing-HPLC, monitored by the fluorescence detection using 2-cyanoacetamide as a post-column labeling reagent. Eight unsaturated disaccharides (DeltaUA-GlcNAc, DeltaUA-GlcNS, DeltaUA-GlcNAc6S, DeltaUA2S-GlcNAc, DeltaUA-GlcNS6S, DeltaUA2S-GlcNS, DeltaUA2S-GlcNAc6S, DeltaUA2S-GlcNS6S) were produced from antler HS by digestion with the mixture of heparin lyases. The total content of 2-O-sulfo disaccharide units in antler HS was higher than that of heparan sulfate from most other animal sources.     

Growth of the first antler in Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>)

In this study, we describe the process of pedicle and first antler growth in Iberian red deer (Cervus elaphus hispanicus) and document relationships among body development, maternal milk supply and composition, and maternal weight on the length of first antlers. Antler length of 53 males of Iberian red deer was measured every 2 weeks from birth to 20 months of age. Deer weight, age, and the date of occurrence of the major events during the antler growth cycle were also recorded. The first evidence of pedicle development occurred when the animals were 38.0 ± 0.6 weeks old and weighed 60.7 ± 0.9 kg. Antler cleaning took place at a mean age of 63.8 ± 0.7 weeks and a mean weight of 91.5 ± 1.8 kg. The antler growth period lasted 16.7 ± 0.4 weeks, and the cleaning period lasted 5.1 ± 0.4 weeks. First antler growth followed a sigmoid curve, reaching a final length of 38.3 ± 1.0 cm. Antler length was positively correlated with body weight during the antler growth cycle. Additionally, the final length of the first antler was related to total milk yield, date of antler growth initiation, body weight at 6 months of age, and the antler growth time interval.     

Deer antler – A novel model for studying organ regeneration in mammals

Deer antler is the only mammalian organ that can fully grow back once lost from its pedicle – the base from which it grows. Therefore, antlers probably offer the most pertinent model for studying organ regeneration in mammals. This paper reviews our current understanding of the mechanisms underlying regeneration of antlers, and provides insights into the possible use for human regenerative medicine. Based on the definition, antler renewal belongs to a special type of regeneration termed epimorphic. However, histological examination failed to detect dedifferentiation of any cell type on the pedicle stump and the formation of a blastema, which are hallmark features of classic epimorphic regeneration. Instead, antler regeneration is achieved through the recruitment, proliferation and differentiation of the single cell type in the pedicle periosteum (PP). The PP cells are the direct derivatives of cells resident in the antlerogenic periosteum (AP), a tissue that exists in prepubertal deer calves and can induce ectopic antler formation when transplanted elsewhere on the deer body. Both the AP and PP cells express key embryonic stem cell markers and can be induced to differentiate into multiple cell lineages in vitro and, therefore, they are termed antler stem cells, and antler regeneration is a stem cell-based epimorphic regeneration. Comparisons between the healing process on the stumps from an amputated mouse limb and early regeneration of antlers suggest that the stump of a mouse limb cannot regenerate because of the limited potential of periosteal cells in long bones to proliferate. If we can impart a greater potential of these periosteal cells to proliferate, we might at least be able to partially regenerate limbs lost from humans. Taken together, a greater understanding of the mechanisms that regulate the regeneration of antlers may provide a valuable insight to aid the field of regenerative medicine.This article is part of a Directed Issue entitled: Regenerative Medicine: the challenge of translation.     

Identification of differentially expressed genes in the developing antler of red deer Cervus elaphus

Understanding the molecular mechanisms underlying bone development is a fundamental and fascinating problem in developmental biology, with significant medical implications. Here, we have identified the expression patterns for 36 genes that were characteristic or dominant in the consecutive cell differentiation zones (mesenchyme, precartilage, cartilage) of the tip section of the developing velvet antler of red deer Cervus elaphus. Two major functional groups of these genes clearly outlined: six genes linked to high metabolic demand and other five to tumor biology. Our study demonstrates the advantages of the antler as a source of mesenchymal markers, for distinguishing precartilage and cartilage by different gene expression patterns and for identifying genes involved in the robust bone development, a striking feature of the growing antler. Putative roles for “antler” genes that encode α-tropomyosine (tpm1), transgelin (tagln), annexin 2 (anxa2), phosphatidylethanolamine-binding protein (pebp) and apolipoprotein D (apoD) in intense but still controlled tissue proliferation are discussed.     

The use of antlers to monitor temporal variation in environmental lead levels: a case study from an industrialized area in Germany

In order to assess temporal changes in ambient lead levels, Pb concentrations were determined in roe deer (Capreolus capreolus) antlers (n=116) that had been collected in the industrialized area of Siegen (western Germany) in the period 1948–2000. Lead concentrations ranged between 0.3 and 166.3 mg/kg dry weight. An overall decline in antler Pb concentration occurred from the mid-1970s onward, and lead concentrations in the samples from the periods 1980–1989 and 1990–2000 were significantly lower than those in samples from previous periods. Our findings indicate a marked decline in lead exposure of the roe deer inhabiting the study area in the recent past. This decline is attributed to a decrease in atmospheric lead deposition, caused mainly by the phase-out of leaded gasoline, and, in addition, by a reduction of lead emissions from stationary sources. The results of the present study and of previous investigations demonstrate that analysis of antler samples obtained in an area over a certain period can be used to reconstruct temporal changes in environmental lead levels.     

Antler growth rate in yearling Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>)

The aims of this study were to describe first antler growth rate and the factors affecting it. Growth rate reached a peak in week 14. Mean first antler growth rate during the whole growing period was 1.95 ± 0.05 cm/week. Lactation influenced antler growth rate as this was positively affected by total protein yield, while antlers grew at a lower rate the later they started to grow.     

Effects of environmental conditions, human activity, reproduction, antler cycle and grouping on fecal glucocorticoids of free-ranging Pampas deer stags (Ozotoceros bezoarticus bezoarticus)

In this study, a commercial enzyme immunoassay (EIA) was validated in detecting glucocorticoids in Pampas deer feces, in order to investigate the influence of several factors on the adrenocortical function. Fecal samples, behavioral data and information concerning male grouping and antlers status were collected at a monthly basis during a 1 year period from free-ranging stags living at Emas National Park, Brazil (18 degrees S/52 degrees W). The results revealed that concentrations of fecal glucocorticoids in winter were significantly higher than those corresponding to spring and summer. In addition, dry season data presented higher levels than during the wet season. Significant difference was found between fecal levels of breeding stags in summer and nonbreeding stags, whereas no difference was observed between breeding stags in winter and nonbreeding stags. On the other hand, males from areas with frequent human disturbance exhibited higher glucocorticoid concentrations and flight distances than individuals from areas of lower human activity. Males with antlers in velvet had elevated levels compared with animals in hard antler or antler casting. Also, we found that glucocorticoid levels were higher in groups with three or more males than in groups with only one male. The flight distances showed positive correlation with fecal glucocorticoid. These data indicate that fecal glucocorticoid provides a useful approach in the evaluation of physiological effects of environment, inter-individuals relationship and human-induced stressors on free-ranging Pampas deer stags.