Trabecula

right|thumb|200x200px|Inside of a bone showing the trabecular structure

thumb|A typical [[carcinoid tumor of the lung showing a trabecular pattern of elongated groups of cells.]]

A trabecula (: trabeculae, from Latin for 'small beam') is a small, often microscopic, tissue element in the form of a small beam, strut or rod that supports or anchors a framework of parts within a body or organ. A trabecula generally has a mechanical function, and is usually composed of dense collagenous tissue (such as the trabecula of the spleen). It can be composed of other material such as muscle and bone. In the heart, muscles form trabeculae carneae and septomarginal trabeculae, and the left atrial appendage has a tubular trabeculated structure.

Cancellous bone is formed from groupings of trabeculated bone tissue. In cross section, trabeculae of a cancellous bone can look like septa, but in three dimensions they are topologically distinct, with trabeculae being roughly rod or pillar-shaped and septa being sheet-like.

When crossing fluid-filled spaces, trabeculae may offer the function of resisting tension (as in the penis, see for example trabeculae of corpora cavernosa and trabeculae of corpus spongiosum) or providing a cell filter (as in the trabecular meshwork of the eye).

Bone trabecula

Structure

Trabecular bone, also called cancellous bone, is porous bone composed of trabeculated bone tissue. It can be found at the ends of long bones like the femur, where the bone is actually not solid but is full of holes connected by thin rods and plates of bone tissue. The holes (the volume not directly occupied by bone trabecula) is the intertrabecular space, and is occupied by red bone marrow, where all the blood cells are made, as well as fibrous tissue. Even though trabecular bone contains a lot of intertrabecular space, its spatial complexity contributes the maximal strength with minimum mass. It is noted that the form and structure of trabecular bone are organized to optimally resist loads imposed by functional activities, like jumping, running and squatting. And according to Wolff's law, proposed in 1892, the external shape and internal architecture of bone are determined by external stresses acting on it. The internal structure of the trabecular bone firstly undergoes adaptive changes along stress direction and then the external shape of cortical bone undergoes secondary changes. Finally bone structure becomes thicker and denser to resist external loading.

Because of the increased occurrence of total joint replacement and its impact on bone remodeling, understanding the stress-related and adaptive process of trabecular bone has become a central concern for bone physiologists. To understand the role of trabecular bone in age-related bone structure and in the design for bone-implant systems, it is important to study the mechanical properties of trabecular bone as a function of variables such as anatomic site, bone density, and age related issues. Mechanical factors including modulus, uniaxial strength, and fatigue properties must be taken into account.

Typically, the porosity percent of trabecular bone is in the range 75–95% and the density ranges from 0.2 to 0.8 g/cm³. It is noted that the porosity can reduce the strength of the bone, but also reduce its weight. The porosity and the manner that porosity is structured affect the strength of material. Thus, the micro structure of trabecular bone is typically oriented and <nowiki></nowiki>grain<nowiki></nowiki> of porosity is aligned in a direction at which mechanical stiffness and strength are greatest. Because of the microstructural directionality, the mechanical properties of trabecular bone are highly anisotropic. The range of Young's modulus for trabecular bone is 800 to 14,000 MPa and the strength of failure is 1 to 100 MPa.

As mentioned above, the mechanical properties of trabecular bone are very sensitive to apparent density. The relationship between modulus of trabecular bone and its apparent density was demonstrated by Carter and Hayes in 1976. The resulting equation states:

<big><math> E = a + b\cdot\rho^c </math></big>

where <math>E</math> represents the modulus of trabecular bone in any loading direction, <math>\rho</math> represents the apparent density, and <math>a,</math> <math>b,</math> and <math>c</math> are constants depending on the architecture of tissue.

Using scanning electron microscopy, it was found that the variation in trabecular architecture with different anatomic sites lead to different modulus. To understand structure-anisotropy and material property relations, one must correlate the measured mechanical properties of anisotropic trabecular specimens with the stereological descriptions of their architecture. Loss of bone mass is defined by the World Health Organization as osteopenia if bone mineral density (BMD) is one standard deviation below mean BMD in young adults, and is defined as osteoporosis if it is more than 2.5 standard deviations below the mean. A low bone density greatly increases risk for stress fracture, which can occur without warning. The resulting low-impact fractures from osteoporosis most commonly occur in the upper femur, which consists of 25-50% trabecular bone depending on the region, in the vertebrae, which are about 90% trabecular, or in the wrist.

When trabecular bone volume decreases, its original plate-and-rod structure is disturbed; plate-like structures are converted to rod-like structures and pre-existing rod-like structures thin until they disconnect and resorb into the body. With osteoarthritis, the underlying bone plays a significant role in cartilage degradation. Thus any trabecular degradation can significantly affect stress distribution and adversely affect the cartilage in question.

Due to its strong effect on overall bone strength, there is currently strong speculation that analysis in patterns of trabeculae degradation may be useful in the near future in tracking the progression of osteoporosis.

Birds

The hollow design of bird bones is multifunctional. It establishes high specific strength and supplements open airways to accommodate the skeletal pneumaticity common to many birds. The specific strength and resistance to buckling is optimized through a bone design that combines a thin, hard shell that encases a spongy core of trabeculae. The allometry of the trabeculae allows the skeleton to tolerate loads without significantly increasing the bone mass. The red-tailed hawk optimizes its weight with a repeating pattern of V-shaped struts that give the bones the necessary lightweight and stiff characteristics. The inner network of trabeculae shifts mass away from the neutral axis, which ultimately increases the resistance to buckling. Besides the difference in distribution, the aspect ratio of the individual struts was higher in woodpeckers than in other birds of similar size such as the Eurasian hoopoe The woodpeckers' trabeculae are more plate-like while the hawk and lark have rod-like structures networked through their bones. The decrease in strain on the woodpecker's brain has been attributed to the higher quantity of thicker plate-like struts packed more closely together than the hawk or hoopoe or the lark.

Trabecula in other organisms

The larger the animal, the higher the load forces on its bones. Trabecular bone increases stiffness by increasing the amount of bone per unit volume or by altering the geometry and arrangement of individual trabeculae as body size and bone loading increases. Trabecular bone scales allometrically, reorganizing the bones' internal structure to increase the ability of the skeleton to sustain loads experienced by the trabeculae. Furthermore, scaling of trabecular geometry can moderate trabecular strain. Load acts as a stimulus to the trabecular, changing its geometry so as to sustain or mitigate strain loads. By using finite element modelling, a study tested four different species under an equal apparent stress (σapp) to show that trabecular scaling in animals alters the strain within the trabecular. It was observed that the strain within trabeculae from each species varied with the geometry of the trabeculae. From a scale of tens of micrometers, which is approximately the size of osteocytes, the figure below shows that thicker trabeculae exhibited less strain. The relative frequency distributions of element strain experienced by each species shows a higher elastic moduli of the trabeculae as the species size increases.

Additionally, trabeculae in larger animals are thicker, further apart, and less densely connected than those in smaller animals. Intra-trabecular osteon can commonly be found in thick trabeculae of larger animals, as well as thinner trabeculae of smaller animals such as cheetah and lemurs. The osteons play a role in the diffusion of nutrients and waste products of osteocytes by regulating the distance between osteocytes and bone surface to approximately 230 μm.

Due to an increased reduction of blood oxygen saturation, animals with high metabolic demands tend to have a lower trabecular thickness (Tb.Th) because they require increased vascular perfusion of trabeculae. The vascularization by tunneling osteons changes the trabecular geometry from solid to tube-like, increasing bending stiffness for individual trabeculae and sustaining blood supply to deep tissue osteocytes.

Bone volume fraction (BV/TV) was found to be relatively constant for the variety of animal sizes tested. Larger animals did not show a significantly larger mass per unit volume of trabecular bone. This may be due to an adaptation which reduces the physiological cost of producing, maintaining, and moving tissue. However, BV/TV showed significant positive scaling in avian femoral condyles. Larger birds present decreased flight habits due to avian BV/TV allometry. The flightless kiwi, weighing only 1–2&nbsp;kg, had the greatest BV/TV of the birds tested in the study. This shows that trabecular bone geometry is related to 'prevailing mechanical conditions', so the differences in trabecular geometry in the femoral head and condyle could be attributed to different loading environments of coxofemoral and femorotibial joints.

The woodpecker's ability to resist repetitive head impact is correlated with its unique micro/nano-hierarchical composite structures.