Z. Kahn-jetter et al., Compressive/tensile stresses and lignified cells as resistance components in joints between cladodes of Opuntia laevis (Cactaceae), INT J PL SC, 161(3), 2000, pp. 447-462
The Cactaceae are a diverse group of plants with a wide variety of morpholo
gies. Many species of Opuntia have segmented stems in which terminal cladod
es may be separated from main-stem cladodes with varying amounts of resista
nce. From a geometric approach, derivations were used to calculate normal (
axial and bending) and shear (transverse force and torque) stresses at join
ts due to the weight of the cladodes. Normal and shear stresses act perpend
icular and parallel to (along) the cross sections of joints, respectively.
Normal stress caused by bending was >10 times that of the mean value of any
other stress. Analyses were performed to determine the relationship betwee
n maximum normal stress and the amount of lignified xylem cells. Such cells
had thicker cell walls compared with the various other cells of stem joint
s that had thin cell walls and that thus would provide the most resistance
to normal stresses. An analogy was made between cactus joints and a composi
te beam with reinforcing rods. In such joints, thin-walled parenchyma cells
might be analogous to concrete that has little resistance to tensile stres
s, while the thick-walled, lignified xylem cells would be analogous to rein
forcing rods. There were statistically significant relationships between no
rmal stresses (from bending and axial loads) and mean percentage of lignifi
ed xylem cells (r = 0.73) and between normal stresses and total areas of li
gnified xylem cells (r = 0.65) (more stress, more reinforcing xylem cells).
Tensile portions of cactus joints had 23% lignified xylem cells, while com
pressive portions had only 10% lignified xylem cells in joint areas (more t
ension, more reinforcing xylem cells). In addition, tensile joint tissues h
ad two to three times more thick-walled, lignified xylem cells in the outer
30% of the radius compared with other joint tissues types (more reinforcin
g near the surface). To our knowledge, this is the first publication to pre
sent mechanical stresses at stem joints of cacti and the first to relate th
ese stresses to characteristics of resisting tissues in the joints of a cac
tus.