Professional Documents
Culture Documents
ix
Acknowledgements
To my dear husband, Giuseppe, and children, Elettra Don’t be trapped by dogma – which is living with
and Jago, for the time that I have stolen from them to the results of other people’s thinking. Don’t let the
write this book. noise of others’ opinions drown out your own inner
To my father for inspiring me to look beyond classi- voice. And most important, have the courage to
cal anatomy, and for an enduring fascination with the follow your heart and intuition.
fascial system. Steve Jobs, Stanford Report, 2005
To Warren Hammer and his assistant, Martha Cook
Hammer, for their suggestions for improving this book
and for helping to make it clear.
x
1
Connective Tissues
Composition of the the metabolic properties of the tissue, the fibres the
mechanical properties, and the ECM the plasticity and
Connective Tissues malleability of the tissue. The most common cell types
Connective tissue (CT) is one of the four major classes are fibroblasts, which produce collagen fibres and oth-
of tissue (the others being epithelial, muscle and nerve er intercellular materials. Cells such as adipocytes and
tissue). It maintains the form of the body and its or- undifferentiated mesenchymal cells are also present.
gans, and provides cohesion and structural support for The proportions of these three components vary from
the tissues and organs. CT derives its name from its one part of the body to another depending on the lo-
function in connecting or binding cells and tissues. It cal structural requirements. In some areas, the CT is
is ubiquitous in the body and can be considered the loosely organized and highly cellular; in others, its fi-
‘glue’ that holds the body parts together. brous components predominate; and in still others,
CT has three main components: cells, fibres and ex- the ground substance may be its most conspicuous
tracellular matrix (ECM) (Fig. 1.1). The cells provide feature. The consistency of the ECM is highly variable
Fibroblasts
Adipocytes
Cells
Macrophages and mast cells
(metabolism)
Undifferentiated
mesenchyme cells
Chondroblasts/ chrondrocytes,
Osteoblasts/ osteocytes
Collagen
CONNECTIVE Fibres
TISSUE (mechanics)
Elastin
Ground
substance Glycosaminoglycan
(viscosity and
plasticity)
Water
Ions
1
and ranges from gelatin like to a more rigid material. proportions. It is clear, colourless and viscous. The fi-
Consequently, the CT ranges in consistency from the bres are of different types, but the principal ones are
gel-like softness of loose CT to the hardness of bone. collagen and elastin and these define the mechanical
The anatomical classification of the various types of CT properties of the tissue.
is based largely upon the relative abundance and ar-
rangement of these components. For example, strong GROUND SUBSTANCE
CTs, such as tendons and ligaments, require a greater
proportion of collagen fibres and fewer cells. Whereas, Ground substance is an amorphous gel-like substance
surrounding the cells. It does not include fibres (e.g.
Connective Tissues
2
are also found in cellular membranes and inside cells, cell receptors and interacting with the cytoskeleton it
and mediate the interactions of the cells and ground confers motility to the cells.
substance. HA is particularly plentiful during embryogenesis
and in tissues undergoing rapid growth, and is present
Hyaluronan
wherever repair and regeneration occur. Depending
Hyaluronan (HA) is the GAG most represented in loose on its chain length, especially when it becomes frag-
CT and is the only one that has no sulfate groups. It mented, HA has recently been shown to have a wide
differs from the typical GAG because it is extremely range of opposing biological functions: such as becom-
Connective Tissues
long and rigid: consisting of a chain of several thou- ing angiogenic, inflammatory and immunostimulatory.
sand sugars, as compared to several hundred or less The turnover of HA is about 2–4 days compared to
found in other GAGs. It also does not bind to a core 7–10 days for the other sulfated GAGs. This means that
protein to become part of a proteoglycan; instead, pro- the HA cells must remain active, otherwise there is the
teoglycans indirectly bind to HA via special linker pro- risk of reduction in the quantity of ground substance.
teins to form giant macromolecules. These hydrophilic The residual products of the GAGs have a feedback
macromolecules are particularly abundant in cartilage effect on the cells and this controls synthesis. It has
ground substance and are responsible for the turgor been established that mechanical distortion of CT cells
pressure that gives cartilage its shape. HA provides represents a stimulus for ECM synthesis (Adhikari et al
the structure as well as turgor to the aqueous of the 2011).
eye and protects fetal vessels from compression in the
Wharton’s jelly of the umbilical cord (Fig. 1.2).
HA provides moisture for the skin by means of its Link proteins
large volume of solvent water (as much as 10,000 Link proteins stabilize the aggregates of proteoglycans
times the volume of the original material). It also pro- in the ground substance and form large bottle-brush-
vides a lubricant for muscles and tendons as they glide like configurations. Of the link proteins, the most well
over skeletal or under aponeurotic fasciae. It is likely known are vinculin, spectrin and actomyosin. They
that these gliding interactions are influenced by the represent the constituents that mediate the interac-
composition and efficacy of the HA-rich ECM. This HA- tions among cells, fibres and other components of
rich layer also protects muscles and supports recovery the matrix and their primary tasks are the binding of
from injury, and stimulates satellite cell proliferation collagen fibres to the cell membrane and the organi-
following loss of muscle fibres. Changes in this HA- zation of the elastic fibres in the ECM. Other specific
rich matrix contribute to pain, inflammation and loss functions for individual link proteins are to guide mo-
of function. HA is abundant during the earliest stag- bile cells through the CT, to control the activity of the
es of wound healing and functions to open up tissue cell nucleus, mitochondria, and Golgi apparatus, and
spaces through which cells can travel. By binding to to connect the cytoskeleton with the ECM. During the
Artery
Artery
Mesenchymal
connective tissue
(or Wharton's jelly)
Vein
FIGURE 1.2 Histology of the umbilical cord, Alcian blue, enlargement 50×. Note how much the mesenchymal CT stains blue, highlighting the
abundance of hyaluronan in the ECM of the umbilical cord.
3
aging process the quantity of link proteins increases and the muscularis externa of the alimentary canal) are
and this reduces the mobility of the CT. able to secrete all of the CT fibre types.
Single collagen fibres usually align along the main
FIBRES lines of a mechanical load. Under pathological circum-
stances, due to changes in the density of ground sub-
There are two types of fibres secreted by CT cells:
stance, the collagen fibres get closer to each other and
collagen fibres and elastic fibres. The abundance and
may form pathological cross-links. This may prevent
preponderance of different types of fibres varies ac-
the ability of a normal collagen network to develop.
cording to the type of CT. Both types of fibre are formed
Connective Tissues
4
Loose connective
tissue
Endomysium
Connective Tissues
Muscular fibre
Perimysium
A Loose connective
tissue
Loose connective
tissue
Nerve fascicle
Perineurium
Epineurium
Adventitia
Loose connective
tissue
C
FIGURE 1.3 Immunohistochemical stain to show the presence and localization of collagen type III in a muscle (A), a peripheral nerve (B) and small
vessels (C). Note the abundance of collagen type III fibres in the endomysium and perimysium, in the perineurium and in the wall of the small
arteries and veins.
and form an elastin matrix. The entire matrix is not form elastic fibres. Instead, the elastin is laid down
engaged during the stretch and recoil of elastic in fenestrated sheets or lamellae arranged in concen-
tissue. tric layers between layers of smooth muscle.
• Fibrillin is a fibrillar glycoprotein. In developing
elastic tissue, it appears before the elastin, and is Connective Tissue Cells
believed to serve as an organizing structure. Many different kinds of cells can be found in CT. The
In most cases, elastic fibres are produced by fibrob- more important cells are the fibroblasts, but also adi-
lasts; however, elastic artery fibres are produced by the pocytes and undifferentiated mesenchymal cells can be
smooth muscle cells of the tunica media. The elastic found. If the adipocytes are numerous and organized
material produced by the smooth muscle cells does into lobules, the CT is referred to as adipose tissue.
not contain fibrillin, only elastin, and as a result does The fibroblasts can differentiate into cells responsible
5
Femoral condyles
covered by cartilage
Connective Tissues
Cruciate ligament
Meniscus
Patella
FIGURE 1.4 Macroscopic aspect of the knee cartilage: femur condyles and patellar surface. The cartilage forms a smooth surface. Note the
cartilage degeneration at the patellar articular surface.
6
Abbott et al (2013) theorize that CTs, especial-
CLINICAL PEARL 1.3 BENEFICIAL RESULTS ly fibroblasts, are part of a whole body cell-to-cell
FROM MECHANICAL LOAD ON FIBROBLASTS communication-signalling network. They state that fi-
AND CIRCULATION broblasts exhibit active cytoskeletal responses within
Mechanical load strongly influences the activity of the minutes of tissue lengthening:
fibroblasts and the deposition of collagen fibres. After a Analogous cell-to-cell signalling involving calcium
sprain or another trauma of the locomotor system, new and/or ATP may exist within CT and may be
collagen fibres will be produced; however, if the patient accompanied by active tissue contraction or
Connective Tissues
is immobilized the collagen fibres will have an irregular relaxation. One can envisage a whole-body web
disposition. This will cause restricted movement and of CT involved in a dynamic, body-wide pattern of
prolong recovery time. Only early movement permits the cellular activity fluctuating over seconds to minutes
correct formation of collagen fibres along the functional reflecting all externally and internally generated
lines of force. mechanical forces acting upon the body.
Loghmani and Warden (2009) injured bilaterally A particular type of fibroblast called the myofibrob-
the medial collateral ligaments (MCL) of 51 rodents last is found in tendons, fasciae and scars (Hinz et al
and used instrument-assisted, cross-fibre massage on 2012). These cells have in their cytoplasm actin fibres
the contralateral ligaments of 31 rodents one week allowing them to contract. During wound repair it is
following injury. They cross-friction massaged the necessary for fibroblasts to convert into myofibroblasts
injured area three sessions per week for one minute per that then create extracellular collagen fibre deposition.
session. Treatment was introduced unilaterally with the They express a smooth muscle type actin–myosin com-
contralateral, injured MCL serving as an internal control plex that closes wounds and speeds up their repair by
(nontreated). Results showed that the treated ligaments contracting the edges of the wound. Upon resolution
were 43.1% stronger (P < 0.05), 39.7% stiffer (P < 0.01), of the injury, these myofibroblasts undergo apoptosis
and could absorb 57.1% more energy before failure (programmed cell death). In wounds that fail to re-
(P < 0.05) than the contralateral, injured, nontreated solve and become keloids or hypertrophic scars, my-
ligaments at four weeks postinjury. On histological and ofibroblasts may persist, rather than disappearing by
scanning electron microscopy assessment, the treated apoptosis. According to Schleip (Schleip et al 2006),
ligaments appeared to have improved collagen fibre these cells also have an important role in determining
bundle formation and orientation within the scar region the basal tone of CT.
when compared with nontreated ligaments.
In a similar study, Loghmani and Warden (2013) used
cross-friction massage on injured MCLs and found that ADIPOCYTES/FAT CELLS
there was not only a temporary increase in vasodilation
Adipocytes can be present in many types of CT as iso-
within the ligament, but an alteration of microvascular
lated cells or as small aggregates. They form a special-
morphology in the vicinity of the healing knee ligaments,
ized CT, called adipose tissue, when they become the
including a larger proportion of blood vessels in the
predominant cell and their main function is the stor-
diameter range of arterioles. These changes persisted for
age of energy as fat. Although the lineage of adipocytes
one week after final intervention.
is still unclear, preadipocytes are undifferentiated fi-
broblasts originating from mesenchymal stem cells
that when stimulated form adipocytes.
We can distinguish two types of adipose cells:
tissue. A scar is collagen deposited by fibroblasts dur-
ing repair.
• Unilocular adipocytes: these are large cells (their
diameter varies from 50 to 100 µ) characterized
Tendons that undergo high rates of stretching may by the presence of a large lipid droplet
be more susceptible to inflammation and eventual de- surrounded by a layer of cytoplasm. The nucleus
generation due to the stretching of fibroblasts. Cyclic is flattened and located on the periphery. A typical
stretching of fibroblasts, and especially increasing the fat cell is 0.1 mm in diameter although some are
frequency of the stretching, increases the production twice that size and others half that size. The fat
of proinflammatory cyclooxygenase enzyme (COX-1, stored is in a semi-liquid state, and is composed
COX-2) and prostaglandin-E2 (Yang et al 2005). There- primarily of triglycerides and cholesteryl esters.
fore, overstimulation of fibroblasts may be responsible These fat cells secrete many proteins such
for repetitive-motion problems. Recent studies (Kaux as resistin, adiponectin and leptin, and they
et al 2013) have shown how eccentric exercises may can synthesize estrogens from androgens.
be more beneficial than concentric exercises regard- Their number can increase in childhood and
ing the rehabilitation of muscles and tendons. There is adolescence, while it remains constant in
reason to believe that the effect of the load pattern of adulthood. When fat cells have increased in size,
eccentric exercise creates greater stimulation of fibrob- about fourfold, they begin to divide, increasing
lasts, which increases collagen synthesis and thereby the absolute number of fat cells present. After
stimulates the healing of the injured tissue. Stretching marked weight loss the number of fat cells does
also causes an increase in tendon fibroblast alignment. not decrease, but rather they contain less fat.
7
Approximately 10% of fat cells are renewed
Loose Connective Tissue
annually at all adult ages. These adipocytes can
organize to form the white adipose tissue (WAT). Loose CT (or areolar tissue) is the most widespread
• Multilocular cells: these are little cells
characterized by the presence of numerous
CT of the body. It is characterized by an abundance of
ground substance, plus thin and relatively few fibres
smaller lipid droplets in the cytoplasm and also and cells (Fig. 1.7). The main cellular elements are
contain a large quantity of mitochondria. These fibroblasts and a smaller amount of adipocytes. Fat
fat cells can organize to form the brown adipose cells are a normal constituent of loose CT, but when
Connective Tissues
tissue (BAT). they are abundant and organized into large lobules
for storage purposes the tissue is better classified
MULTIPOTENT STROMAL CELLS as adipose tissue. The adipocytes present in loose
CT are generally isolated cells or small aggregations
These cells retain the potential of embryonic mesen-
that do not function as storage depots, and their
chymal cells with the capability of differentiating into
principle function is to facilitate gliding and to act
a variety cell types, including osteoblasts, chondro-
as interstitial filler. The adipocytes of the loose CT
cytes, adipocytes, myocytes and neurons. They have a
usually do not increase in volume when individuals
great capacity for self-renewal while maintaining their
gain weight. Collagen is the principal fibre of loose
multipotency.
CT and is arrayed in all directions to form a loose
network in the intercellular material. Many elastic
Classification of fibres are also present.
The loose CT has a viscous, gel-like consistency and
Connective Tissue its consistency may fluctuate in different parts of the
There are three subtypes of CTs: specialized, proper body due to variations in temperature or pH. This CT
and embryonic. The specialized CT includes adipose allows gliding between the various muscles and organs
tissue, bone and cartilage. Regarding the specialized (Figs 1.8 and 1.9) and permits the diffusion of oxygen/
CT, we will be discussing only the adipose tissue due nutrients from small vessels to the cells and the dif-
to its strong relationship to the superficial fascia. Please fusion of metabolites back to the vessels. It is the ini-
refer to other textbooks for information on bone and tial site where antigens, bacteria and other agents that
cartilage. have breached an epithelial surface can be destroyed.
Proper CT is a very large group of tissues comprising It also forms a mesh-like tissue with a fluid matrix
both loose and dense CT (Fig. 1.5). It encompasses all that supports the epithelia, such as the skin and other
organs and body cavities and connects one part with membranes. This CT fills the spaces between various
another. Equally important, it separates one group of organs and thus holds them in place while cushioning
cells from another. All fasciae have been classified as and protecting them; it also surrounds and supports
proper CT, but some authors exclude either the loose the blood vessels.
or the dense from this definition. The embryonic CT A particular type of loose CT is the reticular tissue
includes the mesenchyme (Fig. 1.6) and mucous CT that contains only reticular fibres made of type III col-
(Fig. 1.2). In the following discussion we will describe lagen. The reticular cells have a stellate shape and long
the pertinent features of the proper CT so that we can processes that make contact with neighbouring cells,
correctly classify fasciae. and the subsequent tissue supports a number of bodily
Ligaments
8
Skin
Hypodermis
Connective Tissues
Muscular fibres
FIGURE 1.6 Histological section of the abdominal wall of an embryo of 19 weeks, hematoxylin and eosin stain, enlargement 50×. Note the high
cellularity and absence of well-defined fascial planes.
Collagen fibres
Elastic fibres
Adipocytes
Ground substance
FIGURE 1.7 Histological section of the loose CT, hematoxylin and eosin stain, enlargement 50×. Note the absence of any type of tissue organization.
The collagen and elastic fibres are disposed in many directions and some adipocytes are present.
structures, such as the liver, spleen, bone marrow and hormones that are involved in overall energy home-
lymphatic organs. ostasis. There are different types of adipose tissue, for
example: adipose tissue is commonly distinguished
in WAT and BAT (Smorlesi et al 2012). The WAT is di-
Adipose Tissue vided into two major types according to localization:
Adipose tissue is not merely a tissue designed to subcutaneous white adipose tissue (SWAT) and viscer-
passively store excess carbon in the form of triacylg- al white adipose tissue (VWAT). Sbarbati et al (2010)
lycerols. Mature adipocytes synthesize and secrete used the structural and ultrastructural features of the
numerous enzymes, growth factors, cytokines and adipose cells to classify the WAT into three different
9
Muscle lifted
(enveloped by its
epimysial fascia)
Fat lobules
Loose connective
Connective Tissues
tissue
Muscle enveloped
by its epimysial
fascia
FIGURE 1.8 Macroscopic aspect of the loose CT between pectoralis major and minor muscles. The loose CT creates a gliding surface between the
two muscles and permits their independent contraction. The white lines are the collagen fibres. The empty space between them is occupied in the
living by water linked to the GAGs.
Flexor hallucis
longus muscle
Deep fascia
Loose connective
tissue between the
superficial and deep
compartment
Epimysial fascia
(or epimysium)
of the soleus muscle
Soleus muscle
FIGURE 1.9 Transverse section of the middle third of the leg. The fascia of the flexor hallucis longus is detached from the underlying muscle while
the soleus is strongly adhered to its own fascia. Tractioning the soleus muscle distally simulates a contraction and its fascia follows the muscle.
Note the loose CT between the fascia of the soleus and the flexor hallucis longus. This creates a gliding surface between the two fasciae allowing
an independent contraction and/or passive stretch of the two muscles.
types: deposit WAT (dWAT), structural WAT (sWAT) and cell volume. There are differing microscopic features
fibrous WAT (fWAT). of subcutaneous adipose tissue in different parts of
the body. In the subcutis of the abdomen, the fat cells
WHITE ADIPOSE TISSUE (WAT) are tightly packed and linked by a weak network of
WAT is the major form of adipose tissue in mammals isolated collagen fibres. These collagen fibres are very
(commonly referred to as ‘fat’). It is composed of adi- meager with large cells and few blood vessels (Fig.
pocytes held together by a loose CT that is highly vas- 1.10). In the SWAT of the limbs, the stroma is fairly well
cularized and innervated. White adipocytes are round- represented with adequate vascularity and its cells are
ed cells that contain a single large fat droplet that wrapped by a basket of collagen fibres (Figs 1.11 and
occupies over 90% of the cell volume and the mito- 1.12). In the foot and in areas where a severe mechani-
chondria and nucleus are squeezed into the remaining cal stress can occur, the SWAT has a significant fibrous
10
Umbilicus
Connective Tissues
Subcutaneous
fat tissue of
the abdomen
Skin of the
thorax
Skin of
the hip
Fat lobules
Subcutaneous
hypertrophic
vessel
B
FIGURE 1.10 (A) Macroscopic aspect of the adipose tissue from the abdomen of a fatty cadaver. (B) Adipose tissue of the abdomen removed
from an obese person. Note the large lobules of fat and the paucity of fibrous tissue support. The vessels cross the tissue perpendicularly and
are hypertrophic.
11
Connective Tissues
Dermis
Fat lobules
Fibrous septa
(retinaculum cutis)
Subcutaneous
vessel
Deep fascia
FIGURE 1.11 Macroscopic aspect of the thigh adipose tissue of a standard cadaver. The fat lobules are small and the supporting fibrous tissue is
well represented. The vessels are small and numerous and are sufficient to vascularize the adipose tissue in a homogeneous manner.
12
Fat cells
Vessels
Connective Tissues
Connective
tissue binding
the fat lobules
FIGURE 1.12 Histology of the adipose tissue of the thigh. The adipocytes are small and form regular lobules. Each lobule is supported by CT.
The vessels are numerous. Hematoxylin and eosin stain, enlargement 25×.
Fat lobules
Fibrous
septa Vessels
Skin
FIGURE 1.13 Macroscopic aspect of the fat pad of the heel. In the weight bearing areas the fibrous component (with a white appearance) of the
adipose tissue (with a yellow appearance) increases with each adipocyte having its own thick fibrous shell. This type of fat functions as a cushion
and does not increase in thickness when a person gains weight.
component containing adipocytes with thick distinct adipokines, such as inflammatory cytokines, comple-
fibrous shells (Fig. 1.13). We can recognize a differ- ment-like factors, chemokines, and acute phase pro-
ent composition of the SWAT based on its relationship teins. Its endocrine function includes participating in
with the superficial fascia. The WAT between the skin the regulation of appetite, glucose and lipid metabo-
and the superficial fascia is true fatty tissue and usu- lism, the inflammatory process and reproductive func-
ally increases when a person fattens, while the WAT tions. Subcutaneous and visceral adipocytes originate
between the superficial fascia and the deep fascia is from different progenitor cells that exhibit different
usually looser and generally does not increase in thick- genetic expression patterns. SWAT, compared to VWAT,
ness (see Chapter 2) (Fig. 1.14). responds better to the antilipolytic effects of insulin
VWAT is composed of several adipose depots, in- and other hormones, secretes more adiponectin and
cluding mesenteric, epididymal and perirenal depots less inflammatory cytokines, and is differentially affect-
(Fig. 1.15). VWAT tissue is associated with insulin ed by molecules involved in signal transduction as well
resistance, diabetes mellitus, dyslipidaemia, hyperten- as drugs. According to Sudi et al (2000), the quantity
sion, atherosclerosis, hepatic steatosis, and overall of subcutaneous adipose tissue from the upper body
mortality. is significantly and positively correlated to the level of
The primary function of the WAT is to store energy leptin, suggesting that leptin is under the control of
and act as a cushion. However, it also plays impor- certain subcutaneous adipose tissue depots from the
tant roles as an endocrine/immune organ by secreting upper body.
13
Epidermis
Dermis
Connective Tissues
Superficial fascia
Loose connective
tissue in the deep
adipose tissue
Deep fascia
FIGURE 1.14 Full-thickness histology of the subcutaneous tissue of the thigh, Mallory–Azan stain, enlargement 16×. It is evident that the white
adipose tissue between the skin and the superficial fascia is true fatty tissue, and is formed by adipose lobules surrounded by fibrous septa.
Between the superficial and deep fascia the tissue is looser and the fibrous septa are scarce.
Abdominal
muscles
pulled back
Great omentum
Small intestine
Descending colon
Intra-abdominal fat
(mesenteric depot)
FIGURE 1.15 Macroscopic aspect of the visceral white adipose tissue of the abdomen. The greater omentum was lifted to show the small intestine
with the fat all around the loops. The fat lobules are large and scantily vascularized. The fibrous CT, with its function of support, is almost absent.
14
BROWN ADIPOSE TISSUE (BAT) contain either collagen or elastic protein fibres; there-
The BAT is a specialized adipose tissue and its primary fore, there are dense collagenous CTs and dense elastic
function is thermogenesis. BAT is so-called because types. The collagenous types are far more abundant
it is darkly pigmented due to the high density of mi- and are called fibrous or ‘white’ CT. Elastic fibres, on
tochondria rich in cytochromes. It specializes in the the other hand, appear yellow in unstained tissues
production of heat (adaptive thermogenesis) and li- and are commonly referred to as ‘yellow’ CT (e.g. the
pid oxidation and is especially abundant in newborns yellow ligaments of the spine). Fibroblasts are the only
and in hibernating mammals, but could be found also cells visible and are arranged in rows between the
Connective Tissues
in human adults in the neck, around the aorta and in fibres. Their function is to create the collagen fibres
the supraclavicular, mediastinal and interscapular ar- of the tissue.
eas. BAT contains more capillaries than white fat since The main roles of dense CT are to transmit forces
it has a greater need for oxygen than most tissues. over a distance and to connect different organs/mus-
Brown adipocytes are smaller in overall size compared cles. Collagen fibres are disposed along the direction
to white adipocytes and are polygonal in shape and of mechanical loads present in that specific tissue. The
contain numerous large mitochondria. Whereas, white capacity of dense CT to resist mechanical stress is di-
adipocytes contain a single large fat droplet, brown rectly related to the structural organization of the ECM
adipocytes contain several small lipid droplets. Brown and above all, the collagen fibres.
adipocytes are molecularly Thermogenin + (UCP1+) The dense CT is subclassified as follows:
and leptin–. BAT is highly vascularized and contains a
very high density of noradrenergic nerve fibres. BAT is • Dense, irregular CT has irregularly arranged
collagen fibres and usually comprises the dermis
essential for classical nonshivering thermogenesis; this and fasciae. In the last few years, it has been
phenomenon does not exist when functional BAT is demonstrated that the irregular appearance
absent. In addition, it is essential for cold acclimation- of deep fasciae may be due to its multilayered
recruited, norepinephrine-induced thermogenesis. structure, but in actuality each layer presents its
Heat production from BAT is activated whenever the own regularity (see Chapter 3). Consequently,
organism is in need of extra heat, e.g. postnatally, dur- the deep fasciae could be classified as dense
ing entry into a febrile state, and during arousal from regular CT.
hibernation. Feeding also results in activation of BAT.
A series of diets, apparently all characterized by being • Dense, regular CT is a white, flexible tissue that
contains tightly packed bundles of collagen
low in protein, would result in a leptin-dependent re-
cruitment of the tissue. When the tissue is active, high fibres. All of these fibres run in one direction and
amounts of lipids and glucose are combusted in the are arranged parallel to the direction of forces
tissue. The development of BAT, with its characteristic exerted on the particular body part where the
protein, UCP1 was probably essential for the evolu- tissue is located. This arrangement is typical of
tionary success of mammals, as its thermogenesis en- tendons and ligaments, but according to recent
hances neonatal survival and allows for active life even studies (Benetazzo et al 2011) the deep fasciae
in cold surroundings. Ito et al (1991) demonstrated could also be classified in this group. Purslow
that human brown fat cells begin to show a transfor- (2010) demonstrated that the epimysium and
mation into white fat cells at the infantile stage. This perimysium have a very specific organization
change occurs from the peripheral towards the central that also may classify them as dense, regular CT,
portion of the lobule, so that various functioning cell and Huijing and Baan (2003) demonstrated this
types remain only in the central area of the lobules. In tissue’s role in force transmission (see Chapter 3).
contrast to humans, in hibernating animals the white Finally, a specific discussion of the endomysium
fat cells never occur within the brown fat tissue. Ac- is necessary as it is not clear whether this has
cording with Bartness et al (2010), the BAT presents a structure similar to a microtendon (Purslow
both a sympathetic innervation and a sensory inner- 2010) or if it is better classified as loose CT
vation, which probably has the role of perceiving the (Testut 1905). In the following chapters, the
temperature changes and monitoring the lipolysis. It characteristics and functions of the deep fasciae,
has also been demonstrated that in some BAT depots, epimysium and perimysium will be described in
a parasympathetic innervation exists. greater detail.
We can further classify the regular CT according to
Dense Connective Tissue its function:
Dense CT is characterized by large, robust collagen
fibres that provide a considerable amount of strength
• Dense CT connecting two bones: referred to as
ligaments as they are composed of collagen fibres
to this tissue. Fibres are so numerous that the key positioned parallel to each other and rich in
identifying trait of this tissue is the absence of open elastic fibres (Fig. 1.16).
spaces between cells or fibres. Since the protein fi-
bres are the dominant component of these tissues,
• Dense CT connecting muscles to bones:
referred to as common tendons as they are
the types of fibres and their orientation within these also characterized by collagen fibres positioned
tissues is the basis for the naming scheme. Dense CTs parallel to each other (Figs 1.17 and 1.18), and
15
Radius
Brachioradialis
Connective Tissues
muscle
Interosseous
membrane
Ulna
Flexor ulnaris
carpi tendon
Pronator quadratus
muscle
Wrist joint
FIGURE 1.16 Dissection of the anterior region of the forearm (in a supine position). The interosseous membrane of forearm appears as a broad
ligament joining radius and ulna bones with well evident collagen fibrous bundles. The interosseous membrane divides the forearm into anterior
and posterior compartments, serves as a site of attachment for muscles of the forearm and transfers forces from the radius, to the ulna, and to
the humerus.
16
Muscular fibres
of triceps
Connective Tissues
Calcaneal tendon
Epitenon
Loose connective
tissue
Paratenon
FIGURE 1.17 Macroscopic view of a transverse section of the distal third of the leg showing the calcaneal tendon. The crural fascia splits around
the tendon to form the paratenon. The paratenon envelops both the calcaneal tendon and Kager’s fat pad. At this level some muscular fibres of
the triceps are present. The tendon is formed by closely packed, parallel collagen fibres.
Little vessel
Endotenon
FIGURE 1.18 Histology of the tendon. The fibres are typically arranged in bundles all parallel to each other, bound by the endotenon and rich in
hyaluronan. The blood vessels penetrate into the tendon following the septa of the endotenon. The only cell type present is the fibroblasts and
they are scarce. Only small amounts of ground substance are present. Mallory–Azan stain, enlargement 50×.
17
Small fat lobules
Collagen fibre
bundles with a
Connective Tissues
transverse
orientation
Collagen fibre
bundles with a
longitudinal
orientation
FIGURE 1.19 Macroscopic view of the fascia lata. The fascia lata is a dense CT formed by packed collagen fibre bundles with different orientations.
Loose connective
tissue
Collagen fibre
bundles all parallel
to each other
FIGURE 1.20 Histology of the lacertus fibrosus (myofascial expansion of the biceps brachii muscle into the medial portion of the antebrachial
fascia). The collagen fibres are all positioned parallel to each other and form a fibrous layer. Mallory–Azan stain, enlargement 50×.
18
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Subcutaneous adipose tissue classification. Eur. J.
Abbott, R.D., Koptiuch, C., Iatridis, J.C., Howe, A.K., Histochem. 54 (4), e48.
Badger, G.J., Langevin, H.M., 2013. Stress and matrix- Schleip, R., Naylor, I.L., Ursu, D., et al., 2006. Passive
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J. Cell. Physiol. 228 (1), 50–57. contractility of intramuscular connective tissue. Med.
Adhikari, A.S., Chai, J., Dunn, A.R., 2011. Mechanical Hypotheses 66 (1), 66–71.
load induces a 100-fold increase in the rate of Smorlesi, A., Frontini, A., Giordano, A., Cinti, S., 2012.
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collagen proteolysis by MMP-1. J. Am. Chem. Soc. 133 The adipose organ: white–brown adipocyte plasticity
(6), 1686–1689. and metabolic inflammation. Obes. Rev. Suppl. 2,
Bartness, T.J., Vaughan, C.H., Song, C.K., 2010. 83–96.
Sympathetic and sensory innervation of brown Standring, S., 2008. Gray’s Anatomy, fortieth ed.
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S36–S42.
Sudi, K.M., Gallistl, S., Tafeit, E., Möller, R., Borkenstein,
Benetazzo, L., Bizzego, A., De Caro, R., Frigo, G.,
M.H., 2000. The relationship between different
Guidolin, D., Stecco, C., 2011. 3D reconstruction of
subcutaneous adipose tissue layers, fat mass and
the crural and thoracolumbar fasciae. Surg. Radiol.
leptin in obese children and adolescents. J. Pediatr.
Anat. 33 (10), 855–862.
Endocrinol. Metab. 13 (5), 505–512.
Carano, A., Siciliani, G., 1996. Effects of continuous and
Testut, J.L., Jacob, O., 1905. Précis
intermittent forces on human fibroblasts in vitro. Eur.
d’anatomietopographique avec applications medico-
J. Orthod. 18 (1), 19–26.
chirurgicales, vol. III. Gaston Doinet Cie, Paris, p. 302.
Clayton, H.A., Cressman, E.K., Henriques, D.Y., 2013.
Yang, G., Im, H.J., Wang, J.H., 2005. Repetitive
Proprioceptive sensitivity in Ehlers–Danlos syndrome
mechanical stretching modulates IL-1beta induced
patients. Exp. Brain Res. 230 (3), 311–321.
COX-2, MMP-1 expression, and PGE2 production in
Hinz, B., Phan, S.H., Thannickal, V.J., et al., 2012. Recent human patellar tendon fibroblasts. Gene 19 (363),
developments in myofibroblast biology: paradigms 166–172.
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transmission: muscle relative position and length Benjamin, M., 2009. The fascia of the limbs and back – a
determine agonist and synergist muscle force. J. Appl. review. J. Anat. 214 (1), 1–18.
Physiol. 94 (3), 1092–1107. Cannon, B., Nedergaard, J., 2004. Brown adipose tissue:
Hukinsa, D.W. L., Aspden, R.M., 1985. Composition and function and physiological significance. Physiol. Rev.
properties of connective tissues. Trends Biochem. Sci. 84 (1), 277–359.
10 (7), 260–264. Gao, Y., Kostrominova, T.Y., Faulkner, J.A., Wineman,
Ito, T., Tanuma, Y., Yamada, M., Yamamoto, M., 1991. A.S., 2008. Age-related changes in the mechanical
Morphological studies on brown adipose tissue in the properties of the epimysium in skeletal muscles of
bat and in humans of various ages. Arch. Histol. Cytol. rats. J. Biomech. 41 (2), 465–469.
54 (1), 1–39. Gil, A., Olza, J., Gil-Campos, M., Gomez-Llorente, C.,
Jaroszyk, F., Marzec, E., 1993. Dielectric properties of Aguilera, C.M., 2011. Is adipose tissue metabolically
BAT collagen in the temperature range of thermal different at different sites? Int. J. Pediatr. Obes. Suppl.
denaturation. Ber. Bunsenges Phys. Chem. 97 (7), 1, 13–20.
868–871. Himms-Hagen, J., 1995. Role of brown adipose tissue
Kaux, J.F., Drion, P., Libertiaux, V., et al., 2013. Eccentric thermogenesis in control of thermoregulatory
training improves tendon biomechanical properties: a feeding in rats: A new hypothesis that links
rat model. J. Orthop. Res. 31 (1), 119–124. thermostatic and glucostatic hypotheses for control
Loghmani, M.T., Warden, S.J., 2009. Instrument-assisted of food intake. Proc. Soc. Exp. Biol. Med. 208 (2),
cross-fiber massage accelerates knee ligament 159–169.
healing. J. Orthop. Sports Phys. Ther. 39 (7), Huijing, P.A., 2009. Epimuscularmyofascial force
506–514. transmission: A historical review and implications for
Loghmani, M.T., Warden, S.J., 2013. Instrument-assisted new research. J. Biomech. 42 (1), 9–21.
cross fiber massage increases tissue perfusion and Huijing, P.A., Jaspers, R.T., 2005. Adaptation of muscle
alters microvascular morphology in the vicinity of size and myofascial force transmission: a review and
healing knee ligaments. Complement. Altern. Med. 28 some new experimental results. Scand. J. Med. Sci.
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Purslow, P.P., 2010. Muscle fascia and force transmission. Huijing, P.A., Van De Langenberg, R.W., Meesters, J.J.,
J. Bodywork Mov. Ther. 14 (4), 411–417. Baan, G.C., 2007. Extramuscular myofascial force
19
transmission also occurs between synergistic muscles Rowe, R.W., 1981. Morphology of perimysial and
and antagonistic muscles. J. Electromyogr. Kinesiol. endomysial connective tissue in skeletal muscle.
17 (6), 680–689. Tissue Cell 13 (4), 681–690.
Järvinen, T.A., Józsa, L., Kannus, P., Järvinen, T.L., Sakamoto, Y., 1996. Histological features of
Järvinen, M., 2002. Organization and distribution endomysium, perimysium and epimysium in rat
of intramuscular connective tissue in normal lateral pterygoid muscle. J. Morphol. 227 (1),
and immobilized skeletal muscles. An 113–119.
immunohistochemical, polarization and scanning
Connective Tissues
20
2
Subcutaneous Tissue and
Superficial Fascia
21
Superficial adipose
Skin tissue Superficial fascia
Subcutaneous Tissue and Superficial Fascia
Epidermis
Dermis
Superficial
adipose tissue
with the
retinaculum cutis
superficialis
(skin ligaments)
Superficial fascia
Deep adipose
tissue with
the retinaculum
cutis profundus
Deep fascia
Muscle
22
Umbilicus
Deep fascia
of the external
Rectus sheath
Adherence between
superficial and deep
fasciae along the
inguinal ligament
Deep adipose
tissue
Superficial fascia
of the abdomen
Superficial adipose
tissue
FIGURE 2.3 Macroscopic view of the superficial fascia of the abdomen. It is a very fibrous layer referred to as Scarpa’s fascia.
superficial fascia splits to envelop vessels, nerves or fat superficial fascia, they typically often show a large area
cells, and it seems that the fascia is composed of more of attachment, similar to a fan or a cone. In these areas
than one layer. the superficial fascia appears thicker. It is probable that
The features of the subcutaneous tissue vary through- the arrangement of these septa have contributed to the
out the body, in particular the SAT and DAT differ in great variability in the fascial thickness values reported
thickness, form, and disposition of the adipose lobes in the literature.
and fibrous septa. The retinacula cutis superficialis The superficial fascia and the retinacula cutis form a
(or ‘skin ligaments’ in English textbooks) are usually three-dimensional network between the fat lobules of
almost perpendicular (Fig. 2.4). The retinacula cutis the hypodermis, and this network provides a dynamic
profundus is usually more oblique and thinner than anchor of the skin to underlying tissues. This arrange-
the superficial septa, and creates a clear separation of ment permits a flexible and yet resistant mechanism
the superficial fascia from the deep fascia. Where the of transmission of mechanical loads from multidi-
superficial and deep retinacula cutis insert into the rectional forces. According to Li and Ahn (2011) the
23
Skin lifted
Subcutaneous Tissue and Superficial Fascia
superficial and deep retinacula cutis and superficial the plantar surface of the foot where the DAT is ab-
fascia (which they collectively name as ‘subcutaneous sent. In these areas, the superficial fascia adheres to
fascial bands’) could be considered as structural bridg- the deep fascia, and in SAT the skin ligaments are very
es that mechanically link the skin, subcutaneous layer thick and densely packed, strongly connecting the skin
and deeper muscle layer. Their quantity and morpho- with the underlying planes.
logical characteristics vary according to the region of
the body. For example, the area occupied by the reti-
nacula cutis, with respect to the subcutaneous tissue,
Superficial Adipose Tissue
is thicker in the thigh and calf than in the arm. These The SAT is composed of large fat lobules encased be-
areas are unrelated to hypodermis thickness. The thigh tween fibrous septa (Fig. 2.5). The fat lobules are al-
has the highest average number of retinacula cutis, most circular and the septa (retinacula cutis superfi-
while the greatest average retinacula cutis thickness is cialis or skin ligaments) are well defined and generally
seen in the calf. Regional variations determine the dif- orientated perpendicular to the surface, anchoring the
ferences in mobility of the skin with respect to under- dermis to the deeper planes. The fat lobules are or-
lying tissues and may reflect the composite mechanical ganized in single or multiple layers: depending on the
forces experienced by the body part. For example, in fat content and the thickness of the SAT in the subject
the eyelids, penis and scrotum the adipose tissue and (Fig. 2.6).
the retinacula cutis are absent, and so the skin shows The thickness of the SAT is quite uniform throughout
an increased mobility with respect to the underlying the trunk and generally shows less variation by region
planes. Other examples are the palm of the hand and than the DAT. In the extremities, the SAT is thicker in
24
Skin ligaments (retinacula cutis Superficial
superficialis) Skin adipose tissue
Skin
Superficial fascia
Deep adipose
tissue
Deep fascia
FIGURE 2.6 Macroscopic aspect of the subcutis of the abdomen. Note the fibrous layer (superficial fascia) dividing the hypodermis into two parts:
the SAT and the DAT.
25
the lower limbs than in the upper limbs. The SAT in and nodularity) is more frequent in females. In the
the palm of the hand and the plantar region of the foot elderly, the fat lobules of the SAT are less swollen and
is thin and contains more and stronger vertical reti- the retinacula are less vertical, and thereby connect the
nacula cutis. Therefore, the skin in these areas adheres skin to the superficial fascia with decreased strength.
strongly to the deeper planes. In the dorsum of the Both of these elements are responsible for flaccidity of
hand, a different fascial anatomy permits more move- the superficial tissue with increased age.
ment of the skin with respect to the underlying planes The secretory portion of the sweat glands, hair fol-
because the superficial retinacula cutis is thin. licles and Pacini’s corpuscles are in the SAT. Usually
Subcutaneous Tissue and Superficial Fascia
The SAT thickness varies by subject. In obese sub- these structures are near the retinacula cutis super-
jects, the SAT of the trunk has a mean thickness of ficialis (skin ligaments) that offer protection from
17.2 mm (range 6–35 mm), and in normal-weight sub- stretching and mechanical loads.
jects the mean thickness is 3.7 mm (range 1–10 mm).
In obese subjects, the SAT thickness increases signifi-
cantly and progressively from T10 to the femoral head, Superficial Fascia (Fascia
and in slim subjects the SAT is uniform. The thickness
of the retinacula cutis superficialis also changes by re- Superficialis)
gion of the body and subject. For example, in the trunk The superficial fascia is a fibrous layer of connective
the retinacula cutis superficialis is thicker and stronger tissue, formed by loosely packed interwoven collagen
in the back, giving the SAT of the dorsum a greater re- fibres mixed with abundant elastic fibres (Fig. 2.7). It is
sistance compared to the SAT of the abdomen. Sterzi homologous with the cutaneous muscle layer (pannic-
(1910) found that the skin ligaments of a labourer’s ulus carnosus) found in many lower mammals, where
hand presented a double or triple thickness compared a thin sheet of striated muscle could be found within
to the septa of a sedentary subject’s hand. Sterzi also or just beneath the superficial fascia, and serving to
describes sexual differences in the SAT: females have produce local movement of the skin. For example, a
more fat cells in the SAT, the skin ligaments are thin- grazing animal may twitch the panniculus carnosus to
ner and the fat lobules are arranged in multiple layers. frustrate the attempts of a bird to perch on its back.
These features probably explain why cellulite (hernia- This muscular layer is rare in humans and when found
tion of subcutaneous fat within fibrous connective tis- assumes a precise muscular structural arrangement
sue that manifests topographically as skin dimpling primarily in the neck (platysma muscle) (Fig. 2.8), in
Fat lobules
embedded inside
the superficial
fascia
Fibrous component
of the superficial
fascia
Fat lobules
embedded
inside the
superficial fascia
FIGURE 2.7 Macroscopic aspect of the superficial fascia of the back. The fibrous component (white) and the fat lobules intermingle in the
superficial fascia.
26
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DANCE ON STILTS AT THE GIRLS’ UNYAGO, NIUCHI
I see increasing reason to believe that the view formed some time
back as to the origin of the Makonde bush is the correct one. I have
no doubt that it is not a natural product, but the result of human
occupation. Those parts of the high country where man—as a very
slight amount of practice enables the eye to perceive at once—has not
yet penetrated with axe and hoe, are still occupied by a splendid
timber forest quite able to sustain a comparison with our mixed
forests in Germany. But wherever man has once built his hut or tilled
his field, this horrible bush springs up. Every phase of this process
may be seen in the course of a couple of hours’ walk along the main
road. From the bush to right or left, one hears the sound of the axe—
not from one spot only, but from several directions at once. A few
steps further on, we can see what is taking place. The brush has been
cut down and piled up in heaps to the height of a yard or more,
between which the trunks of the large trees stand up like the last
pillars of a magnificent ruined building. These, too, present a
melancholy spectacle: the destructive Makonde have ringed them—
cut a broad strip of bark all round to ensure their dying off—and also
piled up pyramids of brush round them. Father and son, mother and
son-in-law, are chopping away perseveringly in the background—too
busy, almost, to look round at the white stranger, who usually excites
so much interest. If you pass by the same place a week later, the piles
of brushwood have disappeared and a thick layer of ashes has taken
the place of the green forest. The large trees stretch their
smouldering trunks and branches in dumb accusation to heaven—if
they have not already fallen and been more or less reduced to ashes,
perhaps only showing as a white stripe on the dark ground.
This work of destruction is carried out by the Makonde alike on the
virgin forest and on the bush which has sprung up on sites already
cultivated and deserted. In the second case they are saved the trouble
of burning the large trees, these being entirely absent in the
secondary bush.
After burning this piece of forest ground and loosening it with the
hoe, the native sows his corn and plants his vegetables. All over the
country, he goes in for bed-culture, which requires, and, in fact,
receives, the most careful attention. Weeds are nowhere tolerated in
the south of German East Africa. The crops may fail on the plains,
where droughts are frequent, but never on the plateau with its
abundant rains and heavy dews. Its fortunate inhabitants even have
the satisfaction of seeing the proud Wayao and Wamakua working
for them as labourers, driven by hunger to serve where they were
accustomed to rule.
But the light, sandy soil is soon exhausted, and would yield no
harvest the second year if cultivated twice running. This fact has
been familiar to the native for ages; consequently he provides in
time, and, while his crop is growing, prepares the next plot with axe
and firebrand. Next year he plants this with his various crops and
lets the first piece lie fallow. For a short time it remains waste and
desolate; then nature steps in to repair the destruction wrought by
man; a thousand new growths spring out of the exhausted soil, and
even the old stumps put forth fresh shoots. Next year the new growth
is up to one’s knees, and in a few years more it is that terrible,
impenetrable bush, which maintains its position till the black
occupier of the land has made the round of all the available sites and
come back to his starting point.
The Makonde are, body and soul, so to speak, one with this bush.
According to my Yao informants, indeed, their name means nothing
else but “bush people.” Their own tradition says that they have been
settled up here for a very long time, but to my surprise they laid great
stress on an original immigration. Their old homes were in the
south-east, near Mikindani and the mouth of the Rovuma, whence
their peaceful forefathers were driven by the continual raids of the
Sakalavas from Madagascar and the warlike Shirazis[47] of the coast,
to take refuge on the almost inaccessible plateau. I have studied
African ethnology for twenty years, but the fact that changes of
population in this apparently quiet and peaceable corner of the earth
could have been occasioned by outside enterprises taking place on
the high seas, was completely new to me. It is, no doubt, however,
correct.
The charming tribal legend of the Makonde—besides informing us
of other interesting matters—explains why they have to live in the
thickest of the bush and a long way from the edge of the plateau,
instead of making their permanent homes beside the purling brooks
and springs of the low country.
“The place where the tribe originated is Mahuta, on the southern
side of the plateau towards the Rovuma, where of old time there was
nothing but thick bush. Out of this bush came a man who never
washed himself or shaved his head, and who ate and drank but little.
He went out and made a human figure from the wood of a tree
growing in the open country, which he took home to his abode in the
bush and there set it upright. In the night this image came to life and
was a woman. The man and woman went down together to the
Rovuma to wash themselves. Here the woman gave birth to a still-
born child. They left that place and passed over the high land into the
valley of the Mbemkuru, where the woman had another child, which
was also born dead. Then they returned to the high bush country of
Mahuta, where the third child was born, which lived and grew up. In
course of time, the couple had many more children, and called
themselves Wamatanda. These were the ancestral stock of the
Makonde, also called Wamakonde,[48] i.e., aborigines. Their
forefather, the man from the bush, gave his children the command to
bury their dead upright, in memory of the mother of their race who
was cut out of wood and awoke to life when standing upright. He also
warned them against settling in the valleys and near large streams,
for sickness and death dwelt there. They were to make it a rule to
have their huts at least an hour’s walk from the nearest watering-
place; then their children would thrive and escape illness.”
The explanation of the name Makonde given by my informants is
somewhat different from that contained in the above legend, which I
extract from a little book (small, but packed with information), by
Pater Adams, entitled Lindi und sein Hinterland. Otherwise, my
results agree exactly with the statements of the legend. Washing?
Hapana—there is no such thing. Why should they do so? As it is, the
supply of water scarcely suffices for cooking and drinking; other
people do not wash, so why should the Makonde distinguish himself
by such needless eccentricity? As for shaving the head, the short,
woolly crop scarcely needs it,[49] so the second ancestral precept is
likewise easy enough to follow. Beyond this, however, there is
nothing ridiculous in the ancestor’s advice. I have obtained from
various local artists a fairly large number of figures carved in wood,
ranging from fifteen to twenty-three inches in height, and
representing women belonging to the great group of the Mavia,
Makonde, and Matambwe tribes. The carving is remarkably well
done and renders the female type with great accuracy, especially the
keloid ornamentation, to be described later on. As to the object and
meaning of their works the sculptors either could or (more probably)
would tell me nothing, and I was forced to content myself with the
scanty information vouchsafed by one man, who said that the figures
were merely intended to represent the nembo—the artificial
deformations of pelele, ear-discs, and keloids. The legend recorded
by Pater Adams places these figures in a new light. They must surely
be more than mere dolls; and we may even venture to assume that
they are—though the majority of present-day Makonde are probably
unaware of the fact—representations of the tribal ancestress.
The references in the legend to the descent from Mahuta to the
Rovuma, and to a journey across the highlands into the Mbekuru
valley, undoubtedly indicate the previous history of the tribe, the
travels of the ancestral pair typifying the migrations of their
descendants. The descent to the neighbouring Rovuma valley, with
its extraordinary fertility and great abundance of game, is intelligible
at a glance—but the crossing of the Lukuledi depression, the ascent
to the Rondo Plateau and the descent to the Mbemkuru, also lie
within the bounds of probability, for all these districts have exactly
the same character as the extreme south. Now, however, comes a
point of especial interest for our bacteriological age. The primitive
Makonde did not enjoy their lives in the marshy river-valleys.
Disease raged among them, and many died. It was only after they
had returned to their original home near Mahuta, that the health
conditions of these people improved. We are very apt to think of the
African as a stupid person whose ignorance of nature is only equalled
by his fear of it, and who looks on all mishaps as caused by evil
spirits and malignant natural powers. It is much more correct to
assume in this case that the people very early learnt to distinguish
districts infested with malaria from those where it is absent.
This knowledge is crystallized in the
ancestral warning against settling in the
valleys and near the great waters, the
dwelling-places of disease and death. At the
same time, for security against the hostile
Mavia south of the Rovuma, it was enacted
that every settlement must be not less than a
certain distance from the southern edge of the
plateau. Such in fact is their mode of life at the
present day. It is not such a bad one, and
certainly they are both safer and more
comfortable than the Makua, the recent
intruders from the south, who have made USUAL METHOD OF
good their footing on the western edge of the CLOSING HUT-DOOR
plateau, extending over a fairly wide belt of
country. Neither Makua nor Makonde show in their dwellings
anything of the size and comeliness of the Yao houses in the plain,
especially at Masasi, Chingulungulu and Zuza’s. Jumbe Chauro, a
Makonde hamlet not far from Newala, on the road to Mahuta, is the
most important settlement of the tribe I have yet seen, and has fairly
spacious huts. But how slovenly is their construction compared with
the palatial residences of the elephant-hunters living in the plain.
The roofs are still more untidy than in the general run of huts during
the dry season, the walls show here and there the scanty beginnings
or the lamentable remains of the mud plastering, and the interior is a
veritable dog-kennel; dirt, dust and disorder everywhere. A few huts
only show any attempt at division into rooms, and this consists
merely of very roughly-made bamboo partitions. In one point alone
have I noticed any indication of progress—in the method of fastening
the door. Houses all over the south are secured in a simple but
ingenious manner. The door consists of a set of stout pieces of wood
or bamboo, tied with bark-string to two cross-pieces, and moving in
two grooves round one of the door-posts, so as to open inwards. If
the owner wishes to leave home, he takes two logs as thick as a man’s
upper arm and about a yard long. One of these is placed obliquely
against the middle of the door from the inside, so as to form an angle
of from 60° to 75° with the ground. He then places the second piece
horizontally across the first, pressing it downward with all his might.
It is kept in place by two strong posts planted in the ground a few
inches inside the door. This fastening is absolutely safe, but of course
cannot be applied to both doors at once, otherwise how could the
owner leave or enter his house? I have not yet succeeded in finding
out how the back door is fastened.