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Consulting Editors
Dr Christer Bergsten DVM, PhD Dr Alberto Brizzi DVM Dr Christoph K W Mülling DVM
Assistant Professor Private Practitioner Veterinary Specialist in Anatomy
Department of Animal Environment Parma, Italy Assistant Professor, Department of
and Health Veterinary Anatomy
Swedish University of Agricultural Freie Universität Berlin
Sciences Berlin, Germany
Skara, Sweden

Contributors
Roberto Acuña Adrián González Sagüés Gordon Jones
Private practitioner Director Director
Florida, Uruguay Técnico del Groupo ANKA Dairy Herd Performance
Cuidado de pezuñas Fair Oaks Dairy Farms, Continental Milk
Uri Bargai Pamplona, Spain Producers
Emeritus of Radiology Wheatfield, Indiana
Koret School of Veterinary Medicine Arie Hamoen USA
Rehovat, Israel Chairman
Working Group for Functional Geoff Laurant
R. Boosemann Conformation Traits President
Dierenkliniek Noord-Nederland World Holstein-Friesian Federation Shoof International Ltd
Veendam, The Netherlands The Netherlands Cambridge, New Zealand

Karl Burgi Ruth M. Hirschberg Cristoph Lischer

Program Director and Instructor Institut für Veterinäranatomie, Professor of Equine Clinical Studies
Dairyland Hoof Care Institute Fachbereich Veterinärmedizin Weipers Centre for Equine Welfare
Baraboo, Wisconsin Freie Universität Berlin Division of Veterinary Companion
USA Berlin, Germany Animal Sciences
University of Glasgow Veterinary
James Ferguson Eugene D. Janzen School
Professor of Surgery Associate Dean Clinical Planning Glasgow, Scotland
Chirurgischen Tierklinik Faculty of Veterinary Medicine UK
Universität Leipzig University of Calgary
Germany Calgary, Alberta
Canada

ix
x Consulting Editors and Contributors

Jakob Malmo Ken Nordlund Mike Socha

Registered Veterinary Specialist in Clinical Professor of Food Animal ZINPRO Corporation
Veterinary Medicine Production Medicine Eden Prairie, Minnesota
Cattle Medicine University of Wisconsin-Madison USA
Maffra Veterinary Centre School of Veterinary Medicine
Maffra, Victoria Madison, Wisconsin Mike Stable
Australia USA Bolton Manor Farm
Little Urswick
Dale Miskimins Garrett R. Oetzel Ulverston, Cumbria
Animal Disease Research and Disease Associate Professor UK
Laboratory Department of Medical Sciences
South Dakota State University School of Veterinary Medicine Bimbo Welker
Brookings, South Dakota Madison, Wisconsin Director and Professor
USA USA Large Animal Clinics
Ohio State University
Carlo Maria Mortellaro Pete Ossent Marysville, Ohio
Professor Professor USA
Department of Veterinary Clinical Department of Veterinary Pathology
Sciences University of Zurich Jan-Willem Wopereis
Division of Surgery Zurich, Switzerland Rector
Faculty of Veterinary Medicine Hulshofstraat 10 7135 JV
University of Milan Greg Penner Harreveld, The Netherlands
Milan, Italy College of Agriculture
University of Saskatchewan Kazuhiro Yoshino
Karen Mortensen Saskatoon, Saskatchewan Akabane Animal Clinic
Rineg, Denmark Canada Akabane
Tahara, Aichi
Kerstin Mueller Randy Shaver Japan
Professor Professor and Extension Dairy
Klinik für Klaurntiere Nutritionist
Freie Universität Berlin Department of Dairy Science
Berlin, Germany College of Agriculture and Life Sciences
University of Wisconsin
Jonathan Naylor Madison, Wisconsin
Professor USA
Large Animal Clinical Sciences
Western College of Veterinary Medicine
52 Campus Drive
Saskatoon, Saskatchewan
Canada
Foreword
When Professor Greenough invited me to write a foreword to this book,
I assumed it to be a new edition of his Lameness in Cattle which had already
reached its third edition in 1997. It was not until the proofs arrived that
I noticed the new title Bovine Laminitis and Lameness and realized that it
was an entirely new text. Book titles do not change by accident, so I took
some moments to explore both the prior books and the pages of this
new text.

Why had laminitis been elevated into a title role? I went to my bookshelf and
found my copy of the first edition of Lameness in Cattle, published in 1972.
I located laminitis under a section entitled Metabolic Diseases and found the
entries ‘acute’ and ‘chronic’ laminitis discussed in less than ten pages. There
are histological photographs of thrombi in vessels of the corium. There is a
picture of a cow with a familiar arched back. However, it is both humorous
and gratifying to read the entire discussion of the prevention of laminitis in
a single sentence: ‘Little may be possible in the way of prevention if the cause is
associated with metritis or retained fetal membranes or a particular intensive
fattening system except where a particular foodstuff, such as wheat bran or oilseed
cake, is incriminated.’ Ten pages on laminitis and one sentence on prevention!
As I turned to this new book, I found over a hundred pages on laminitis and
those pages are dominated by discussions of risk factors for laminitis. Gratified
to realize the profound expansion of knowledge in this area, I concluded
that the elevation of laminitis into the title was appropriate.

The intervening 35 years have also seen the emergence of the ‘herd’ as a
unit of interest by the veterinary profession and this book reflects that
growth. We still characterize a hoof as normal or abnormal based upon the
appearance of lesions, but we now characterize the herd as normal or
abnormal based upon the prevalence of specific problems. In treating the
herd, our interventions are as likely to be focused on housing, rations, foot
bath concentrations, or track management as on the cows themselves.
While herd-level risk factors and preventive interventions are a major focus
of this book, the fundamental core of veterinary practice and this book
remains that of making an informed diagnosis of the individual animal
problem. The terminology, photographs, and discussions of specific lesions
should quickly establish this book as the current standard textbook on
bovine lameness. On reflection, I believe that the quality of discussion that
extends across both specific lesions and herd interventions is perhaps
unique and is the greatest strength of this text.

xi
xii Foreword

Several features make these pages very friendly to readers. In color-coded

blocks within each chapter, you will find a Glossary, Key Concepts, and
Technical Comments that stand out for convenience. Understanding that
most users of this textbook will not read it cover to cover, but will instead
search through individual chapters for answers to specific questions, Professor
Greenough has repeated many of the key concepts in related locations
throughout the book. Because literature citations make for disrupted sentences,
they are not offered within the text itself. Should additional information be
desired, a list of references is provided at the end of each chapter. Finally, the
inclusion of color photographs brings new clarity to the many illustrations
of hoof lesions to be found in these pages.

I would like to close with some personal comments about Professor

Greenough. I have been a veterinarian for 30 years. Within a few months
after entering practice, I purchased a copy of the little green first edition of
Bovine Lameness because it had become self-evident that my training on
lameness in veterinary college had been insufficient. That edition served me
well for several years. In the early 1980s, I recall driving to the Minnesota
Herd Health Programming Conference in St. Paul because Paul Greenough
was scheduled to present a series of lectures. As he finished the presentations,
I stood in a queue of people with questions. I held a handful of photographs
of hoof lesions from one of my client’s herds and sought his opinion. I still
recall how he examined the pictures, sat down to discuss the history of the
herd, and offered pointed advice that opened to me a dawning awareness of
the relationship between ruminal acidosis and the lesions I had photographed.
Other contributions continued to arrive over the subsequent decades. For at
least 40 years, Professor Greenough has occupied a position as the central
authority on bovine lameness in North America, collecting and advancing
our knowledge, serving our profession and the cattle industries, and advocating
for animal welfare. On behalf of the hundreds or perhaps thousands of
veterinarians who have improved their services to cows and their owners
because of his career, I say thank you to Professor Paul Greenough.

Ken Nordlund, DVM

January 2007
 CHAPTER 1

Introduction

GLOSSARY

Lameness: Modification of gait; a clinical sign of many diseases and

disorders.
Disease: A sickness caused by an infectious agent.
Disorder: A sickness resulting from an accident or a failure in body
function.
Risk factor: Any stressful circumstance making an animal more prone
to lameness.
Insult: The point in time when a risk factor negatively affects an animal.

KEY CONCEPTS

‘The veterinary practitioner faced with a herd lameness problem on a farm needs
to examine farm management rather than merely treating a long series of
individual cases’ (Chesterton)

• Lameness is a clinical sign and should not be regarded as a disease or

disorder.

• Risk factors affecting the incidence of lameness also affect the

incidence of other conditions, such as reproductive problems and
mastitis.

• An individual skilled in recognizing lameness is 2.5× more likely to

identify an affected animal than a person who is unskilled.

1
2 1 / Introduction

INTRODUCTION part of the text, providing emphasis and reinforcement

to the verbal message or idea. (See Fig. 1-1.) Many
The objective of this book is to present information in colleagues from around the world have given copies of
language that is easy to understand. English will not be photographs they have taken, to the author. This help is
the mother tongue of every reader and idiomatic speech deeply appreciated. The author wishes to apologize for
varies throughout the English-speaking world. For this a poor memory regarding illustrations that credit
reason a glossary is provided. ‘anonymous’ donors.
The primary target reader is the veterinarian who
practices cattle medicine and surgery. The bulk of the
information falls into the category of ‘need to know.’ ABOUT REFERENCES
Another objective of the book is to make this type of
information understandable to other individuals who In this book, the reader will find very few references in
work with practicing veterinarians, such as animal the text. This follows the trend in contemporary scien-
scientists, claw trimmers, nutritionists, and progressive tific books which omit references from the body of the
producers. There are several exceptions to this need- work. Text that is congested with references is uncom-
to-know policy. Chapters 2 and 4 are dedicated to new fortable to read. An author can no longer accommodate
and specialized information concerning the structure, the contemporary explosion of information.
function, and pathophysiology of the digit. This knowl- A bibliography is provided with most chapters. These
edge is basic to understanding certain disease processes. are included as a starting point for literature searches
In other chapters, ‘Technical Comments’ are clearly for use by research workers and graduate students. The
identified as containing information that is only majority of references have been taken from refereed
necessary for the benefit of students and scientists who journals or textbooks usually found in libraries, together
need a deeper understanding of the scientific principles. with the Proceedings of the International Symposia on
A feature termed ‘Key Concepts’ has been included Disorders of the Ruminant Digit. Where an author has
in order to help the reader identify topics or ideas of written multiple papers on the same subject, the most
greatest importance from a practical point of view. The recent publication is cited.
remainder of the text explains, elaborates, or even repeats
some of these concepts.
The author considers that photographs are an AN HISTORICAL COMMENTARY
important component of the communication process. ON LAMENESS
The captions should, for the most part, be considered as
We tend to think of preventing lameness as a modern
notion, but this is not the case. First-century Romans
appear to have taken good care of their oxen:
‘Cattle will be less likely to go lame if their feet are
washed in plenty of cold water when they are
unyoked after work and if their hocks, the crowns of
their hooves, and the division itself between the two
halves of the hooves are rubbed with stale axle grease.’
Furthermore, Roman veterinarians were among the
earliest researchers to record their lameness-related
activities. Once they had perfected the ‘solea ferria,’ a
metal shoe for horses, they turned their attention to a
similar device for cattle. The problem for the Romans
was to increase the distance of a day’s march before
draft oxen became lame. To this end, they created the
‘demi solea’ (Fig. 1-2).
In 1534, the monk Petrus Magni produced a manual
Figure 1-1 Cows at pasture in Costa Rica. Cattle will lie
on how to run the agricultural business of a large
down for up to 14 hours each day. During this period they
will ruminate and produce 108–308 liters (23–68 imperial monastery. He emphasized the importance of good
gallons or 28–81 US gallons) of saliva each day. Saliva is the bedding and cleanliness to ‘protect cattle hooves from
natural buffer to ruminal acidosis. Encouraging dairy cows to rotting.’ The concern about preventing lameness in
lie down for prolonged periods is one of the keys to cattle is reflected in a book by Hess (1887) who also
controlling subclinical laminitis. describes the destructive influence of manure and
 Animal Welfare 3

partly on the prevalence of this clinical sign and the

economic importance of the causal diseases. In another
sense, lameness may have even greater importance.
Well-established control protocols can be implemented
with respect to both mastitis and reproductive disorders;
however, this is not the case with herd lameness. The
increase in the incidence of lameness, particularly in
dairy herds, is partly due to the fact that production
technologies are being adopted without the user being
aware of the implications to animal health.

ANIMAL WELFARE

The conditions causing lameness are among the most

painful diseases or disorders affecting cattle. Other painful
conditions such as renal colic or displaced abomasums
are not as ubiquitous as lameness. It is the sheer number
of lame cows that makes bovine lameness the important
welfare issue that it is. In some dairy herds, 60% of the
cows can be lame in any given year; 20% of the cows
can be lame at any given time in a problem herd.
It is not known how severe the pain is for the cow.
Figure 1-2 A demi solea is a metal shoe ingeniously fixed Pain in the feet is shown by a cow as changes in her body
with cords. Topical dressings were kept in place by a cloth language and this is used as the basis for scoring the
bag located between the claw and the shoe. The Romans did
severity of lameness (pp. 34 and 35). It is presumed that
not attach shoes with nails; this came some centuries later.
(Courtesy of Mosby, Veterinary Medicine: An Illustrated
cows feel pain in their feet much as we do as there is a
History) dense network of sensory nerve fibers and a high
number of nerve endings, particularly in the sole heel
bulb region.
It is known that the incidence of lameness has
moisture and points out the importance of hygiene and increased considerably in dairy herds during recent years
cleanliness. Other publications from the 19th century and now has reached a point at which it is causing an
concerning bovine lameness are noted in the bibliog- unprecedented level of suffering among cattle. This
raphy, attesting to an early interest in the problem. thought is supported by the considerable amount of fund-
Prior to 1960, there were two books of note dedi- ing dedicated to lameness research. A good example is
cated exclusively to bovine lameness: Malattie degli arti the LameCow 2002–2006 project with a budget of
dei bovini written by Professor Guiseppe Gerosa (first =
C 30,543,890 administered by partners in the EU.
published in Milan in 1929) and Klauenkrankheiten by Fundamental to the issue of animal welfare is the
Hess & Wyssmann (1931). These books reflect the state genetic selection of dairy cows. The genetic potential for
of knowledge at that time, which was quite consid- milk production in Holsteins has doubled in just over
erable. Other excellent books published in the first half 20–30 years. Workers in some countries are alarmed
of the 20th century dealt with anatomy (Wyssmann that selection for production has intensified at the
1902), microanatomy (Mueck 1928) and the ortho- expense of selection for non-production traits such as
pedic surgery of both the horse and the cow (Pfeiffer & ‘feet and legs,’ resistance to disease, and other factors
Williams 1900). that contribute to longevity and functional efficiency.
Pain (even discomfort) causes considerable stress.
Pain is only the tip of the iceberg in that for every pain-
THE CLINICAL IMPORTANCE producing lesion there are likely to be many other
OF LAMENESS lesions in the same or other claws. However, pain may
not be the only factor stressing dairy cattle. It is now
Lameness has come to be considered as the most believed that stressors in the environment, referred to as
important animal health issue next to reproductive ‘management risk factors,’ increase the likelihood of
disorders and mastitis. This evaluation is probably based lameness-causing lesions to develop in the claw. It has
4 1 / Introduction

been shown that these same stresses are implicated in • Value of the time of the animal attendant nursing
the occurrence of other disorders, particularly those the cow
affecting fertility and perhaps mastitis. • Loss of milk due to pain (20% of the production
Cow barns are designed to be convenient for farm for the lactation can be lost if the animal becomes
workers rather than for the comfort of cows. Manage- lame during peak lactation)
ment risk factors have attracted a great deal of attention • Negative effect on reproductive efficiency
over the past 10 years, causing the subject of ‘cow • Loss of milk if antibiotic therapy is used
comfort’ to emerge as a science of real concern to those • 10% of lame animals are culled prematurely;
seeking measures to reduce the incidence of herd lame- the salvage value of the culled animal will only
ness. For example, it is now believed that cattle can partly compensate for the rearing costs of a
suffer psychological stress when a dominant animal heifer which will have a lower milk yield than a
interacts with a submissive animal. It is also thought mature cow
that cows are stressed when they stand for long periods; • Loss of body condition
inactive on hard concrete surfaces. However, correcting • Reduction of the economic life of a cow.
risk factors associated with barn design often presents
insurmountable financial difficulties for dairy producers. There have been numerous estimates of the cost of
An important study undertaken in the UK has reported a single case of lameness. One lame cow will cost the
that dairy farmers who lack knowledge and skill are producer at least US$350. Estimates for losses expe-
those who have herds with the highest incidence of lame- rienced in large high-production herds run into many
ness. Another study found that a skilled observer could thousands of US dollars annually.
identify 2.5× more animals lame that could an individual
who was unskilled. The implication is that many lame
animals fail to be treated. Furthermore, in Australasia, EFFECT OF LAMENESS ON
where cattle are herded along trackways, there is a REPRODUCTIVE EFFICIENCY
noticeably higher incidence of lameness in herds where
attendants are not patient and gentle with the animals. When a cow becomes lame, she loses her status of domi-
In Sweden, the importance of animal welfare in dairy nance among the rest of the herd. She becomes less
herds is already a matter of public concern. In that competitive for resources such as water, forages, and
country, the consumer is willing to pay a premium for concentrates. Lameness also causes the interval between
milk (green milk) produced on dairy farms that have been calving and conception to increase. One of the reasons
certified as meeting specified criteria for animal comfort. for this is that the associated pain reduces the ability of
Toe abscess is a condition affecting yearling beef a lame cow to ride another during oestrus. This in turn
calves and is described on pages 100–102. The cause of will reduce the cowman’s ability to detect heat.
this condition is unknown. However, circumstantial The cow is likely to spend more time lying down and,
evidence suggests that standing for long periods while in so doing, becomes more liable to injury by other cows
being transported may be involved. Current laws in and/or to suffer from bed sores. When lame, a cow
North America only require that cattle shall be unloaded spends more time lying during a single lying event, but
every 48 hours. In order to avoid animals falling down less total time lying in a day. A cow will also stand longer
and being trampled, they are packed tightly into trucks. in a free stall before attempting to lie down and will
Possible movement is so restricted that blood cannot change positions much less often while lying down.
move through the feet and pressure inside the claws is In one study it was reported that lame cows had a
thought to increase, causing discomfort. Toe abscesses lower conception rate at first service (17.5% versus
have been found most frequently 10 days after trans- 42.6%) and a higher incidence of ovarian cysts (25.0%
portation and are so painful that calves must lie down versus 11.1%) than controls. Cows becoming lame
and frequently contract pneumonia and die. within the first 30 days postpartum are associated with
a higher incidence of ovarian cysts, a lower likelihood of
pregnancy, and lower fertility than control cows.
ECONOMIC IMPORTANCE One further sequel of stress may be the slower invo-
lution of the uterus after parturition.
The following factors have to be taken into considera- Bacterial endotoxins originating from the break down
tion in calculating the cost of a single case of lameness of bacterial walls (e.g., caused by ruminal acidosis) have
in a dairy cow: a negative effect on ovarian function. Endotoxins also
• Veterinary fees have a number of direct and indirect local effects in the
• Cost of medication organ systems, including the claw.
 Lameness in Different Production Systems 5

developed by the National German Agriculture Society

TECHNICAL COMMENTS (DFG). The ‘LameCow’ system is available in different
languages including English. Electronic recording is
As stress from pain caused by lameness increases, it is future oriented and has many advantages for cow-side
believed that the secretion of adrenocorticotrophic data-recording. For the time being claw trimmers prefer
hormone (ACTH) will also increase with an associated to write their findings on a record sheet.
reduction in the liberation of luteinizing hormone
(LH). These physiological changes will result in
significant alterations in the manifestations of heat, LAMENESS IN DIFFERENT PRODUCTION
and have shown to result in reduced, aberrant, or SYSTEMS
asynchronous LH release and abnormalities in
ovulation. Lameness is directly related to and influenced by the
system of cattle management.
A fundamental role in the pathogenesis of fertility
disturbances is played by the negative energy balance
(NEB) which is invariably experienced by the lame Intensive Dairy Production
cow. Studies have shown that this causes a reduction
of the secretion of gonadotrophin releasing hormone The greatest increase in the incidence of lameness has
(GnRH) and LH which are essential for starting and taken place in high-production intensively managed
maintaining cyclic ovarian activity. Progesterone dairy herds. Problems have resulted from greater
secretion during the luteal phase is also strongly confinement and increasing herd size. Many cows
reduced in cows experiencing NEB. Prompt treatment spend their entire lives walking on concrete. Restricted
and recovery are essential to reduce the duration of freedom of movement results from poor barn design.
NEB combined with stress. There is an increasing tendency, as herd size increases,
As a consequence of these hormonal disturbances, for feedstuffs to be purchased from different sources.
there is a high probability of pregnancy failure, This results in the quality of the feedstuffs being incon-
anestrus, and, under severe prolonged stress, atrophy sistent and contributing to poor nutritional manage-
of the ovaries. Herds with the highest incidence of ment.
lameness are usually those having the highest There has been a remarkable increase in the preva-
production and are the most intensively managed. lence of digital dermatitis since the disease was first
reported in 1974 (pp. 207–211). There has also been an
increased incidence of subclinical laminitis (pp. 40–49)
and the disorders that complicate this condition.

RECORDING FOOT LESIONS

Pasture Managed Dairy Production
A record of the incidence and prevalence of lesions in (see Chapter 8)
the feet of cattle is essential in order to determine the
risk factors likely to be involved in herd lameness In New Zealand and South West Australia, dairy cows
problems. This is a very important task for claw are maintained on pasture. The predominant cause of
trimmers who should be recording all lesions they lameness is trauma occurring when cattle are herded
encounter during trimming sessions. Some lesions to be from the pasture to the milking station. Until recent
recorded may not have developed to the stage where years, dairy cows have received no concentrate supple-
they are causing lameness or which are being resolved mentation. Under these circumstances, milk production
by appropriate trimming methods. Further information was much lower than in intensively managed herds but
should be contributed by the veterinarian, much of the incidence of lameness was quite low. Today lame-
which will speak to the severity and treatment of the ness is on the increase in herds in which concentrate
lesion. Dairymen must record all cases of lameness, supplements are given.
particularly those caused by infectious diseases treated In various countries in South America, extensive
by farm staff. These various inputs give a comprehensive management of dairy cows is practiced, but supplemen-
picture of the herd lameness problem. tation of the ration with concentrates is common.
There are electronic data recording systems avail- Silage is also fed during the winter months. Under these
able now which have been developed by or in collabora- conditions, the incidence of lameness is much higher
tion with hoof trimmers. Such an electronic system was than it is in Australasia.
6 1 / Introduction

In temperate climates, some herds graze pastures Barkema H W, Westrik J D, Keulen Van K A S et al 1994
only during the summer months. The transition periods The effects of lameness on reproductive performance,
during which cows go to grass for the first time and in milk production and culling in Dutch dairy farms.
the fall when they are housed full time are periods when Preventive Veterinary Medicine 20:249–259
Baumgartner C, Distl O 1990 Genetic and phenotypic
problems can be observed.
relationships of claw disorders and claw measurements
in first lactating German Simmental cows with stayability,
milk production and fertility traits. Proceedings of VI
Cow-Calf Beef Herds International Symposium on Disorders of the Ruminant
Digit, Liverpool, UK p 199
Herds of beef cattle usually are maintained on pasture Bonsma J C 1973 In: Cunha T J, Warwick A C, Koger M (eds)
year round. Problems can arise if the quality of the Factors affecting the calf crop. University of Florida Press,
winter diet is significantly inferior to that during the Miami
spring and summer. Capuco A V, Smith J J, Waldo D R et al 1995 Influence of
On the Canadian prairies the incidence of vertical prepubertal dietary regimen on mammary growth of
Holstein heifers. Journal of Dairy Science 78:2709–2725
fissures is, on average, 17% of adult beef cattle, with
Collick D W, Ward W R, Dobson H 1989 Associations
the incidence in some herds reaching 60%. The same between types of lameness and fertility. The Veterinary
condition has been reported in southern Sweden. The Record 125:103–106
cause of this problem may be related to sudden changes De Stefanis 1854 Metodo curativo della gangrenoso falangeo
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Noble, K M, Trebble J E, Harvey D et al 2000 Ultrasonography 5:99–103
and hormone profiles of persistent ovarian follicle (cysts) Wyssmann E 1902 Anatomie der Klauenlederhaut. Bern
induced with low doses of progesterone in cattle. Journal Universitat Veterinärmed. Fak. Diss
of Reproduction and Fertility 120:361–366
 CHAPTER 2

Microstructure and Function

of the Bovine Claw

GENERAL INTRODUCTION

GLOSSARY

Foot: Strictly speaking, the word foot describes the Keratinization or Differentiation:
region from the hock to the apex of the claw. The unidirectional development of epidermal
In this book the word ‘foot’ will describe the cells (produced in the basal layer) to cornified
digital region distal to the fetlock. horn cells. It is characterized by synthesis of
keratins and intercellular cementing substance:
Claw: The word ‘claw’ is used to define the organ at
it is terminated by transformation of the living
the end of the digit of cattle, i.e. capsule and
epidermis in claw horn, i.e. cornification
contents. The same structure in the horse
(cell death).
is referred to as the ‘hoof.’ The claw comprises
the hard cornified claw capsule plus all the Cornification: The final stage of epidermal
structures enclosed by the capsule – bones, differentiation at which a solid/stable horn cell
joints, ligaments, tendons, fat, cushions, has been formed and which is connected to
connective tissue, blood vessels, and nerves. neighboring cells.
Claw Capsule (Capsula Ungulae): The capsule is the Desmosomes (Macula Adherens): Desmosomes are
shoe-like structure composed of horn into which cell contacts establishing a mechanical stable
the pedal bone and surrounding soft tissue fit. cell-to-cell contact. Each desmosome consists of
It is sometimes referred to as the ‘shoe.’ two halves. Each half belongs to one of two
neighboring cells. Each half desmosome has
Proliferation: The production of new epidermal cells
intracellular and extracellular components.
by division of living cells in the basement layer
(mitosis).

8
 The Epidermis 9

Trying to remember long Latin names is at best difficult

and at worst boring. Furthermore, anatomical termi-
nology can be confusing. Nevertheless, knowledge of
anatomy is essential if some of the complex problems
involved in lameness are to be understood. Sometimes
several terms refer to the same structure. In this chapter
the terminology will be based on the simplest descriptors
that are most widely used in the field today.

THE EPIDERMIS

The pedal bone and related structures are completely

covered with modified skin. This modified skin, as it does
in skin elsewhere in the body, consists of three layers –
subcutis, dermis, and the superficial epidermis. The epidermal
capsule is composed of inner living cells and a thick layer
of dead, cornified cells, the claw horn. This capsule is referred
to as the claw capsule, which is a slipper-shaped struc-
ture. For convenience, anatomists have broken down the
claw capsule, and the underlying dermis and subcutis,
into five component segments. These are the coronary
band (periople segment), the wall (the coronary segment),
the epidermal lamellae (wall segment), the solear segment,
and the bulbar segment. Please note that in the preceding
sentence the correct anatomical term is placed in paren-
Figure 2-1 A section of the sole showing: (D) The dermis
theses following the term that is in current usage. The with dermal papillae which represent the tubules in the sole.
microstructure of each segment has unique characteristics. (E) The epidermis. The arrows indicate the dark staining
However, all of the segments have a ‘living epidermis’ ‘mitotic’ cells of the ‘living’ epidermis. The lighter stained
separated from the dermis by a basement membrane. It cells of the remainder of the living epidermis demonstrate
is at this level many pathological processes are initiated. the progressive loss of their nuclei from top to bottom of the
Beneath the dermis is the subcutis, in some segments picture. (H) This darker layer represents the fully keratinized
modified into cushions. cells of the outer layer of the sole. (Courtesy of C K W Mülling)
See Figures 2-1–2-3.

b c

Figure 2-2 These pictures show the cellular structure

of the claw wall. (a) The top layer is the vascular
dermis, the middle layer is the living epidermis and the
lower half represents the keratinized horn of the sole.
a
(b) This is a cross-section of the keratinized wall showing
the distribution of tubules each plugged with medullary
debris (incomplete, cornified horn cells). (c) The scanning
electron micrograph shows the proximal opening of a
tubule. The papillary pegs of coronary dermis fit into
these openings. (Courtesy of C K W Mülling)
10 2 / Microstructure and Function of the Bovine Claw

THE LIVING EPIDERMIS

TECHNICAL COMMENTS

The epidermal cells are connected permanently to the

basal layer on one side and extend out to the surface
of the wall on the other. This intercellular connection
is made by desmosomes and intercellular cementum.
The latter is the major connection between the horn
cells.
Inside each epidermal cell there are electron-dense
plaques. Connected to these two plaques are keratin
filaments (tonofibrils) which form the ‘cytoskeleton.’
The cytoskeleton is connected to the desmosome.
In the space between the cells, specific desmosomal
glycoproteins establish cell-to-cell adhesion. It is
important to understand that the tonofibrils do not
cross between cells.
The basement membrane (BM) consists of three
layers. The lamina densa forms an electron-dense
midline, on the outside of which is the lamina rara
which makes a foundation for the mitotic cells of the
living epidermis. Both of these layers are produced
by the living epidermal cells. The innermost layer of
the basement membrane is the lamina fibroreticularis
which, as its name implies, consists of anchoring
collagen fibers which connect the basement
membrane to the network of collagen fibers in the
dermis. This layer is produced by dermal fibroblasts
and is the part of the basement membrane that is
susceptible to degradation by matrix
metalloproteinases (MMPs).

The living epidermis sits on the basement membrane

(Fig. 2-4) and it also has different layers of cells. The
first is the stratum basale, with cells dividing mitoti-
cally. The next layer is the stratum spinosum, in the
cytoplasm of which keratin filaments can be observed
radiating from the desmosomes (intercellular adhe-
sions, Fig. 2-5). The keratin filaments form an internal
Figure 2-3 This picture shows the transition across the
three-dimensional cytoskeleton which establishes the
living epidermis from the intensely stained mitotic cells
mechanical strength of the epidermal cell while
approximating the dermis at the top of the picture to
the flattened keratinized, dark staining cells that have retaining flexibility. The third layer, the stratum
lost their nuclei at the bottom. (Courtesy of C K W Mülling) granulosum, exists only in the regions where soft horn
 The Living Epidermis 11

Figure 2-4 Scanning electron micrograph (SEM) showing

a group of basal epidermal cells on the surface of a
dermal lamina. The basal cells have long basal processes Figure 2-5 Desmosomes (arrows) connecting two
reaching into the dermis. In the dermis under the basal neighboring epidermal cells in the stratum spinosum.
cells a capillary is opened containing red blood cells. Bundles of tonofilaments connect the cytoskeleton to the
The dermal tissue between basal cells and the vessel wall intracellular plaques of the hemidesmosomes. (Courtesy of
is the tissue that oxygen and nutrients have to travel through C K W Mülling)
on their way to supply the avascular epidermis. (Courtesy of
C K W Mülling)

is produced, i.e., the bulb and periople. The living for activated MMPs which are enzymes designed to
epidermis is nourished via diffusion from the micro- degrade collagen – the major component of connec-
vasculature of the dermis. tive tissue. The MMPs play an important role in the
The basement membrane is frequently referred to normal turnover of connective tissue. An increase in
as the dermo-epidermal junction. It is of particular the activated form of MMPs is also believed to be
importance in regulating the proliferation and differ- responsible for pathological alterations in some forms
entiation of the mitotic cells of the living epidermis. of laminitis.
The regulatory cascade leading to increased basal Vasoactive substances and endotoxins originating
proliferation is activated by a variety of mechanical in the rumen as the result of acidosis, metabolic
and biochemical stimuli including growth factors disorders, or inflammatory processes, and/or hormones
and cytokines. Growth factors, including epidermal either interfere directly with the microcirculation in the
growth factor (EGF), migrate through the basement claw or trigger the release of cytokines, which in turn
membrane to bind at their specific receptors in the affect the tissue integrity in the claw. The result is a
basement layer (stratum basale). The signaling between disturbance in or even disruption of horn production
dermis and epidermis is sometimes referred to as for short or long periods and with varying degrees of
‘cross-talk.’ severity.
The collagen of the innermost layer of the base- The forgoing suggests that two distinctly different
ment membrane (lamina fibroreticularis) is a substrate pathophysiological events are possible. Collagen degra-
12 2 / Microstructure and Function of the Bovine Claw

dation and horn disruption may occur singly or in

concert.
Pressure on the sole of the claw is transferred via
the epidermis to the basal cell layer, stimulating
proliferation of cells and thereby accelerating the
production of horn. Functional claw trimming aims
to distribute load evenly between both claws. The
objective of trimming is to avoid pressure overload of
one sole leading to increased horn production and
enlargement of a region of the sole, usually the heel
(overburdening). Functional claw trimming interrupts
the vicious circle which occurs in mechanical overload
of the claw. On the other hand thinning of the sole in
routine claw trimming also increases pressure on the
keratogenic cells, stimulating the production of healthy
horn.
In the wall the mechanical stimulus for increased Figure 2-7 An electron micrograph (EM) of the dermo-
proliferation and subsequent horn production is created epidermal junction. The basement membrane (BM) is
by direct tensile force rather than pressure. located between the basal processes of a basal epidermal
See Figures 2-6–2-8. cell and the underlying dermis. Keratin filaments (KF) stabilize
the epidermal cell. They are anchored to hemidesmosomes
in the cell membrane (arrow). Outside the cell membrane
Formation of claw horn follows the electron-lucent ‘lamina rara’ and then the dense
‘lamina densa’. These two are product of the epidermis.
Proliferation basal cells The connection to the dermal collagen system (asterisk
indicates a collagen fiber) is established by anchoring fibers
Differentiation

of the lamina fibroreticularis (arrowheads) which is a product

of the dermal fibroblasts. (Courtesy of C K W Mülling)
stratum
spinosum

Cell death
horn

Vicious circle in mechanical overload

increased
proliferation

increased
increased
rate of horn pressure
formation
Figure 2-8 Low power EM of the dermo-epidermal
junction. (E) Epidermis, (B) Basal cell. Arrows indicate the
position of the basement membrane. (D) Dermis.
increased Asterisk indicates a dermal capillary containing three red blood
thickness
of horn cells, which is in close proximity (1–2µm) to the basement
membrane. The distance between the capillary and the basal
Figure 2-6 Composite illustration of growth and vicious epidermal cell would be the diffusion distance for supply
circle in mechanical overload. (Courtesy of C K W Mülling) with oxygen and nutrients. (Courtesy of C K W Mülling)