Question of the day –
Why does the cut grass grow back, but when you prune a tree limb, it stops
growing from that end?
You cut the ends of the blades of grass and they grow back
in a week or so. But have you ever seen a limb regenerate on a tree? It would
be unusual; if you cut the limb off, the tree forms a knot as the trunk grows
around the cut stub.
The difference lies in the kinds of plants grass and trees
are, and how these types of plants grow. Most trees are dicotyledon (di = two,
and cotyledon = embryonic leaf)
plants, while most grasses are monocotyledons
(mono = one). The differences between
these types of plants are many, but the names come from the various
embryonic forms they take in their seed.
Monocots generally do not overwinter well, are herbaceous (without woody stems), have
fibrous root systems, and have flowers with petals in multiples of three. The
leaves of monocots usually have veins that are parallel to the direction of
growth, and the leaves are most often long and thin. They way they move water
and nutrients (in the xylem and phloem, respectively), is constructed in small
islands, with many bundles of xylem and phloem spread throughout the stem.
Dicot plants, on the other hand, are often perennial plants
that survive winters well. They can herbaceous or arborescent (woody), and their transport systems (phloem and xylem)
are arranged in concentric circles. Long, deep taproots are more common in
dicots, as are flowers with petals in groups of four or five and broad leaves
with networked veins.
Monocots and dicots have similar structures (leaves,
flowers, stems, etc.) but they arrange them differently and they have different
properties. One thing they have in common is that their parts are generated
from similar cells, in structures called meristems.
A meristem (meristos = divisible, and em = the ending from xylem and phloem) is vascular tissue that can divide; they are the source of all new cells in growing plants. Meristem cells act much like stem cells; they each have the potential to become many different types of cells. However, I could not find evidence that the older word, meristem, was the source of the term stem cell.
All plants have meristems, both monocots and dicots - no exceptions, if you can believe it. The smallest flowering plant, Wolffia globosa, has meristems, though the whole plant is only a millimeter long. Some meristem features are found only in dicots and others only in
monocots, one type of meristem that both plant types share is the apical
meristem.
Shoot apical
meristems make new stems, shoots, leaves and other above ground structures.
The pluripotent cells of the meristem divide and differentiate to become the
cells that are needed. The structure of the typical meristem is shown in the
picture to the above.
In grasses, there are shoot apical meristems, although they
may sometimes be called intercalary
meristems. They are located at the base of the leaves and the stem, and
allow for re-growth of the leaf after it is grazed by herbivores…. or mowed
down by Harry Homeowner. Therefore, blame the evolution of low placed meristems
as an adaptation to deal with prehistoric cows and goats for your need to mow
each week.
Node
meristems are forms of intercalary meristems. Growth can
continue
to take place from each node. On the left is bamboo, a
good
example of a monocot grass with nodal meristems.
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On the other hand, most trees are dicots. Their shoot apical
meristems are located at the ends of their limbs and top of the trunk. As the
tree grows, limbs branch off from the trunk, each forming from a different shoot
apical meristem.
The apical meristems at the ends of the limbs and top of the
trunk take the shape of buds in the spring. Each bud will become a short twig
and leaf. In the fall, the leaf falls off, the tree goes dormant, and the next
spring a new bud forms on the end and the process is repeated.
This is how a branch becomes longer. If you prune back a
limb, rarely will it grow from the pruned end – you cut the meristem cells away!
The limb may grow from one of the branch points behind where you pruned, but
not from the cut end.
Trunks get taller and limbs get longer from these apical
meristems, but there are also axillary (lateral) meristems produced when a limb
branches or a leaf grows from the twig. However, most of these meristems do not
support growth; they remain dormant.
The meristem at the top of the trunk and the end of the limb are dominant; they produce plant
hormones to prevent growth from the meristems below them (on the trunk) or closer
to the trunk (on limbs). This is why many trees have a single, central trunk.
If that meristem is lost, one or more may become dominant and several trunks
may form.
Auxin is the hormone that induces and maintains dormancy in
lateral meristems, while cytokinin hormones promote growth from the laterals. For a good review of their roles, see the paper of Muller and Leyser from 2011. Other
plant hormones are relevant for induction and emergence from dormancy induced by winter for
all meristems. Abscisic acid (ABA) induces winter dormancy
while gibberellins break the dormancy. A 2012 study has defined gene expression
patterns during short and long-term cold spells and how they affect the
ABA/gibberellin ratio and how short cold periods alter ABA concentrations.
A new study has investigated how fruits can also induce dormancy in meristems. So much energy is put into fruit development that energy must be stolen from plant growth. The fruits themselves produce auxin to save energy by stopping meristem function. This is why fruit trees have alternating years of fruit vs. growth; when they make fruit they don’t grow, and when they grow, they don’t make much fruit.
In some cases, if the shoot apical meristem is lost, the lateral
meristem on a branch can become the dominant shoot apical meristem and the
branch will begin vertical growth as a new trunk. However, this isn’t always
the case (see picture to the right).
These are the ways trees get taller and fuller, but it's
also related to how they get thicker. Apical meristems give rise to a thin line
of cells that form the procambium
and the secondary meristem all
around the periphery of the trunk or limb.
Just as apical meristems form buds and leaves when the
winter breaks, so do secondary meristems form new vascular tissue all around
the periphery each growing season, this tissue is called the vascular cambium. We see their work as
the rings that can be used to age a tree (dendrochronology). Twigs don’t stay
twigs because each year their secondary meristems make them thicker.
Exception one - some things we call trees are actually
monocots, not dicots. If you look at the wood from a palm tree or a cordyline,
you can see the small, dispersed groupings of vascular tissue that are typical
of monocots. The reason they grow taller and appear more like trees is that
they produce more lignan than other monocots, so they can grow taller. Lignan is the stiffest of the plant
structural tissues and provides the resistance to gravity and wind.
Palm trees also have secondary mersitems, similar to the vascular
cambium in dicot trees that add girth. However, palm secondary meristems just
add more vascular bundles, not the rings of vascular tissue you see in dicot
trees.
The second exception has to do with regrowth of trees even
when the complete limb or trunk is cut away. There are meristems that get
buried by the expanding wood layers of the trunk or when a limb grows thicker.
Regeneration of whole trees by stump sprouting is based on
loss of inhibitory chemical signals and stimulation of stem cells, much like
last week’s discussion of the stem cells in your nail matrix and how they can
regenerate the tips of your fingers. Maybe soon we will catch up to plants and
be able to grow new limbs.
Smith HM, & Samach A (2013). Constraints to obtaining consistent annual yields in perennial tree crops. I: Heavy fruit load dominates over vegetative growth. Plant science : an international journal of experimental plant biology, 207, 158-67 PMID: 23602111
Leida C, Conejero A, Arbona V, Gómez-Cadenas A, Llácer G, Badenes ML, & Ríos G (2012). Chilling-dependent release of seed and bud dormancy in peach associates to common changes in gene expression. PloS one, 7 (5) PMID: 22590512
Müller D, & Leyser O (2011). Auxin, cytokinin and the control of shoot branching. Annals of botany, 107 (7), 1203-12 PMID: 21504914
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