The long and short of it - the story of internodes

The structural units of a higher plant consist of a node,
an internode, and one ore more buds and leaves at the
node.  These units can be repeated indefinitely to form
a shoot, or modified in various ways to alter the
architecture of the plant.

Plant stems are organized into repeated units consisting of nodes and internodes.  Nodes seem to be where all the action is.  Leaves are attached at the nodes, and most often there is also a bud at the base of each leaf that has the potential to develop into a new shoot.  The new shoot also will develop as a series of nodes and internodes.


Internodes are essentially the sections of stem between nodes. Ho humm - end of story.  Not quite!  The internode in fact is a dynamic part the plant, the part responsible for most actual growth in height.  You can see the lengthening of internodes most dramatically in vines, which are adapted to extend themselves as rapidly upward as possible to reach the light zone.  In vines, cell division and expansion continue in the internodes well below the tip, propelling several sets of expanding leaves ahead of them.  Once reaching the light zone, internodes will end growth sooner, resulting in a bushier crown suitable for gathering light.  Tree saplings also show dramatic internode elongation in the shade, and many more humble plants will send their flowers rapidly upward by means of strikingly elongate flower stalks.

Internodes are quite distinct in most trees and shrubs (left), but are extraordinarily
elongate in vines like the Clematis (right).

The bamboo's elongate internodes are the secrets to its
height.

Bamboo shoots contain all the
nodes of a full stem packed close
together.  Expansion of all the
internodes, more-or-less
simultaneously, results in very
rapid growth.

Bamboos are grasses evolutionarily adapted to compete with trees, and they do so with the most extraordinary internodal elongation of all.   Each bamboo shoot begins as a compact bud, the "bamboo shoot" of culinary commerce.  Within that massive bud, the nodes and internodes for the entire shoot (culm) are preformed, but the internodes are extremely short.  When growing conditions are right, the entire bamboo shoot elongates through simultaneous elongation of all of its internodes, reaching full height in a matter of days.  The legendary "Chinese bamboo torture" consisted of strapping a prisoner over a bamboo shoot just beginning its expansion phase.  The sharp point of the shoot tip pressing against the back quickly became horribly painful.

This Pachypodium, in the Apocynaceae,
has virtually no internodes, but grows
slowly through the accumulation of
tissues at the base of each leaf.

This Neoregelia in the Bromeliad, forms a concise rosette
of leaves.

Many plants effectively have no internodes.  They are very short, or slowly build up trunks just from the bulk of the tissues around the leaf bases.  Palms and cycads are prime examples, as are many desert succulents.  Plants that hug the surface of the ground have short stems with leaves crowded into a rounded pattern referred to as a rosette (named for its resemblance to the crowded petals of a rose).  Strawberry plants, bromeliads, venus fly traps, sundews, and African violets are examples.

This tiny Drosera  rosette is dwarfed by its much larger flowers.  The flower
stalks consist of greatly elongate internodes, in contrast to the virtual lack
of internodes within the rosette of leaves.

The stalk of the Egyptian papyrus plant
(Cyperus papyrus) is a single elongate
internode.  The tassle at the top consists
of several dozen nodes packed closely
together through lack of any internode
elongation.

The growth of internodes is controlled by hormones, particularly gibberellin.  Dwarf forms of vegetables or other crops can be created by genetically limiting the plant's ability to produce or respond to gibberellin.  And plants that are normally compact, such as cabbages, can be stretched out like vines by applying the hormone to the young plants.

Gaillardia daisies are normally compact rosettes, but
under the influence of gibberellins can stretch out via
elongate internodes.

Internodes are therefore the unsung heroes of plant growth.  They determine whether a plant will be a skinny, stretched-out vine, a squat mound of leaves, or something in-between.

The first "bamboos"

Earlier ("The grasses that would be trees," March 18, 2012), I described the unique pattern of development that results in the tall, lightweight, and very strong stems of bamboos.  The key to the rapid growth of bamboos is a combination of lightweight, hollow construction, plus a process of growth involving intercalary meristems in each internode that all elongate more-or-less at the same time.  Before there were grasses, before in fact there were any seed plants, a group of spore-bearing relatives of ferns discovered virtually the same growth form.  These were the horsetails, formally known as the Sphenophytes. 

Like bamboos, horsetails send up new shoots from buds
on underground rhizomes.  Each bud contains a complete
compressed stem, with many nodes and internodes packed
closely together. Intercalary meristems within each internode
become active at the same time, adding new tissues to each
and raising the stem rapidly.  The true leaves are modified
into toothed, cup-like structures that protect the tender
growing region of each internode.  From Kerner and
Oliver, The Natural History of Plants, 1904, Fig. 190.

Very few of these sphenophytes survive today, but you can see the bamboo-like form in the stems of modern horsetails.  Like bamboos, the horsetail stem is hollow and its wall fortified with fibers.  Also like bamboos, the young horsetail shoot forms as a condensed bud, with nodes and internodes of the entiren stem crowded together.  A basal intercalary meristem in each indernode begins expansion in coordination with all the others in the shoot, resulting in rapid upward growth.  Leaves at each node are reduced to stiff bracts that protect the tender growing region at the base of the internode. 

Giant horsetails, commonly referred to the
genus Calamites, grew like bamboos and
dominated the coal-forming swamps of
the Carboniferous Period. From Smith,
Cryptogamic Botany, 1955,  Fig. 151.

From the late Devonian, Carboniferous and Permian periods, some 350-300 million years ago, giant tree-like horsetails, growing up to 100 feet high, dominated early forests, sprouting from underground rhizomes, just like modern bamboos. They most likely elongated fairly rapidly, but develeped a modest amount of wood to support their large crown of branches.  

A modern horsetail, growing in a ditch beside the
road in Washington State, is just as at-home in the
21st century as its ancestors were 300 million years
ago.  It continues to compete with neighboring
vegetation through its rapid growth from preformed buds
in the spring.  The true leaves are modified into
 bracts that protect the growing  tissues above each node.
Photosynthesis is conducted by tissues in the main
stem as well as by the whorls of slender stems at each node.

Modern horsetails are for the most part fairly modest in size, living in shaded moist areas alongside the descendents of their other ancient companions, ferns and clubmosses.  The largest, up to 8 ft or more in height, are found oddly in moist streamsides in dry areas of Central and South America.  For an image, click on the link below, or if it is no longer active, do a simple web search for Equisetum giganteum: http://www2.fiu.edu/~chusb001/GiantEquisetum/Images/NorthernChile/LlutaRailroadScale2.html