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Tips for the Municipal Arborist: Root Physiology
edited by Leonard E. Phillips, Jr.
From City Trees, The Journal of The Society of Municipal Arborists
Vol 35, Number 4
July/August 1999
How do roots grow and develop into an extensive network capable of holding
a tree 100 feet in the air, against 50 mile per hour winds? This article
will describe how roots grow and interact with the rest of the tree and
the soils in which the tree is growing. This article will be followed by
an article in the next issue of City Trees about solving root problems
in our urban environments.
The roots of all trees are required for anchorage, absorption of nutrients
and water, and the storage of starch. The development of a root system
is dependent upon the plant genetics, the soils, and the environment. The
first part of this article will describe how roots grow and function.
A tree's root system must balance its shoot system, but not in weight
or dimensions. The root system must be able to supply the shoot system
with sufficient moisture and nutrients so that the shoots can manufacture
enough food to support the root system. This balance will be disturbed
by pruning branches, damaging roots, or attacks by insects or disease.
Roots develop internally rather than from buds as occurs on stems and
the parts of the tree above the ground. To do this, the root has several
parts. They are described as follows from the youngest part of a root to
the oldest part:
Root Cap - This part of the root is at its very tip. The cap
protects the tip of the root as it is forced through the soil by the elongating
tissue behind it. Old cells on the cap will slough off the surface and
lubricate the movement of the root through the soil. These cells are constantly
being replaced.
Apical Maristem - This part of the root provides the cells for
the root cap in front and for the region of elongation behind this area.
Region of Elongation - Active cell division and elongation in
this part of the root forces the cap through the soil against the mass
of the tree.
Region of Root Hairs or Differentiation - It is in this area
that cells develop a more mature form and are differentiated into the epidermis
and cortex. Root hairs are formed in the epidermis or surface layer of
cells. The root hairs serve to absorb water for the plant and live for
only a short period of time. They do not become lateral roots. Firs, redwoods
and Scots Pine do not have root hairs. Instead, they absorb water and nutrients
through the thin walled epidermis. In contrast to this, other trees such
as the Redbud and Honeylocust have root hairs that last for several years.
Within the cortex are the endodermis, pericycle, phloem, and xylem.
Lateral Roots - In the area behind the region of root hairs,
lateral roots are formed. Lateral roots begin by sending out a root cap,
apical meristem tissue, etc. into a new area of soil.
Root Forms
Roots can take three forms, a tap root system, a fibrous root system, or
a combination of both. A tap root grows downward and provides anchor support
for the top growth. Fibrous root systems consist of many lateral roots
which grow horizontally to stabilize a tree. Most species will grow a tap
root as a seedling until a certain distance or obstacle is encountered,
then the root system changes and continues to grow as a fibrous system.
Most root systems of trees grow in the top three feet of the soil. This
is where oxygen and organic matter are most prevalent. The small feeding
roots are in the top six inches especially in a forest or mulched area.
In ideal conditions, a tree's roots will spread out two to three times
the crown diameter or a radius distance equal to the height if the soil
is right for the tree.
Roots need a minimum of 3 percent oxygen in the soil to stay alive.
Twelve percent is needed for new roots to form. Fortunately, most normal
soil is about 20 percent oxygen. However, if surface changes have occurred,
and the soil became compacted and contained only 5 percent oxygen, the
existing roots would survive but new roots could not grow, therefore the
tree would become stressed.
Soil types can also influence root development. While clay soils might
have insufficient oxygen levels at three feet deep, sandy soils could have
15 percent oxygen at 5 feet deep.
Root Microorganizms
Trees grow at their best in forest soils. When trees are brought into the
cities and planted in urban soil, they will not do well. The major cause
of this failure is the lack of microorganisms in the soil. Soil microorganisms
consist of animals such as earth worms, protozoa and nematodes. Earth worms
carry organic matter from the surface deep into the soil while their holes
provide a means of water and oxygen to percolate deep into the soil layers.
Protozoa, the most numerous one celled organisms, are useful in decomposing
organic matter and bacteria in the soil. Nematodes are a microscopic worm
that also decomposes organic matter, although some harmful nematodes will
invade and injure the roots of plants.
Other microorganisms consist of algae, bacteria, and fungi:
Algae assist in dissolving minerals and creating soil.
Bacteria decompose organic matter. Some bacteria are very useful
in compost piles, while other bacteria will thrive in anaerobic composts.
Certain bacteria are also useful in legumes for fixing nitrogen to the
roots while others denitrify nitrates. There are also some harmful bacteria
that cause diseases in trees.
Fungi are the most important microorganism in the soil. Although
many fungi are responsible for causing diseases such as rots and wilts,
they are important because they form a symbiotic relationship with a plant's
roots. This association is called mycorrhizae. The word mycorrhizae comes
from myco meaning fungus and rhiza meaning root. A tree can not survive
without mycorrhizae.
Mycorrhizae
The mycorrhizae association occurs when the fungi grow around a root and
invade the outer layer of root cells. The fungi then act like root hairs
to extract minerals from the soil. In poorer soils, the fungi explore the
soil extensively in search of nutrients more efficiently than root hairs
can. The fungi can also absorb soil moisture more easily and increase a
trees's drought tolerance. Mycorrhizae will not perform efficiently in
highly fertilized soils. Therefore the tree has to rely on less efficient
root hairs, which reduce the effectiveness of the fertilizer.
Mycorrhizae roots not only increase a tree's drought tolerance, they
increase tolerance to soil compaction, high temperatures, toxic materials,
salt, extremes of Ph, and root diseases. Three types of the most common
mycorrhizae are:
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Ectomycorrhizae which are fungi that form a mantle or sheath around
roots with no fungal penetration of root cortical cells.
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Endomycorrhizae which are fungi that penetrate cortical cells of
fleshy young roots but form no mantle or sheath.
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Ectendomycorrhizae are those fungi that both penetrate root cortical
cells and form root-surrounding mantles.
At Arizona State University, researchers are studying a type of endomycorrhizae
called vesicular arbuscular mycorrhizae (VAM). VAM fungi have unique vegetative
and reproductive life stages. During their vegetative stage, they develop
a hyphal matrix (or mycelium) comprised of runner, penetrating and
absorbing
hyphae.
Runner hyphae follow roots as they grow in the soil. Sometimes
runner hyphae grow out into the soil probing for more roots.
Penetrating hyphae branch off runner hyphae and infect fleshy
young roots usually within a short distance of the root growing tip. Once
inside the root, these hyphae pierce root cortical cell walls and form
arbuscules between the root cell wall and cell membrane, called the plasmalemma.
Arbuscule are organs for exchanging nutrients with root cells. In addition
to arbuscules, penetrating hyphae also produce storage organs called vesicles
in spaces between root cells.
Finally, absorbing hyphae branch off of runner hyphae in a fanlike
pattern. They grow into soil surrounding the infected root and absorb water
and nutrients. The absorbed water and nutrients subsequently move to the
penetrating hyphae and then into the infected root.
Phosphorus is mainly insoluble and immobile in soil. VAM fungal hyphae
explore the soil beyond the root zone and acquire phosphorus for roots.
In the past, researchers thought that all VAM fungi, regardless of species
or geographic region, were functionally equivalent. This meant that an
increased uptake of phosphorous was the sole cause of VAM promotion of
plant growth. Based on that assumption, researchers thought a plant's need
to be mycorrhizal was related directly to the amount of phosphorus in the
soil. If soil phosphorus was low, then mycorrhizal fungi were necessary
to facilitate phosphorus extraction. However, if soil phosphorus was adequate
or high, then mycorrhizal fungi became little more than root parasites
levying a carbon tax on the plant.
The concept of mycorrhizae does not mean that a root/fungal association
is always beneficial. Situations occur in which a fungus benefits the plant
under one set of conditions but harms it under another, or the fungus benefits
one plant but harms another. Researchers are learning from this increasingly
complex picture of VAM fungi that it is inadequate to measure the efficiency
of a mycorrhizae solely in terms of host plant phosphorus nutrition or
an enhancement of plant growth. Other adaptational responses may be equally
important. For example, under drought conditions, VAM promotion of growth
should take a back seat to improved plant water relations. Or, plants recently
transplanted into a landscape could benefit more from VAM fungi that promote
root exploration of soil rather than promote shoot growth.
During the 1980's, researchers noticed that some mycorrhizae caused
the roots of herbaceous plants to become thicker, less branched and more
elongated. Recently, plant scientists at Arizona State University discovered
that the VAM fungus, suppressed shoot growth and caused roots of a landscape
tree to become thicker and less branched under drought conditions, compared
with drought stressed trees not infected with VAM. Mycorrhizal suppression
of growth and thickening of roots may be a drought survival mechanism promoting
storage of carbon reserves in roots until drought conditions improve.
Compared with a highly branched or dichotomous root system, a sparsely-branched
or herringbone root system would be more beneficial under conditions of
soil stress or intense root competition because of its ability to explore
a larger soil volume. In addition, mycorrhizal colonization of sparsely
branched roots increases the total root absorbing surface because fungal
hyphae act as root conduit extenders. Such changes in root branching caused
by VAM fungi could have significant ramifications for successful transplant
establishment of landscape trees and shrubs.
Roots attract mycorrhizal hyphae by excluding organic acids that act
as chemical signals. As root tips continue to grow out into the soil, the
infected root segment begins to harden. This hardening causes an end to
the mycorrhizae and ensures that fungal colonization of roots occurs only
on fleshy portions of the root system that are most active in absorbing
water and nutrients.
Root Interaction with Soils
Soils are very often the cause of tree root problems. We try to plant trees
in compacted soils, soils destroyed by construction, or soils subject to
flooding or poor drainage. Soils that have problems with pH, salt, nutrient
levels, toxins, and amendments, cause problems for tree roots. A so]] analysis
should be made prior to planting a tree. A tree can not develop top growth
if it can't develop root growth.
Planting sites should be large enough to accommodate the tree's roots
at maturity. This means 4 square feet of surface area for every inch of
diameter the tree is expected to attain, or 2 cubic feet of soil for every
square foot of the future crown. The soil should also be between 20 and
30 inches deep for maximum vigor. The more surface area the better. Sites
with pavement and barriers to healthy root development should be avoided.
Proper planting techniques are essential for tree root survival.
If the soil is typical of an urban site - concrete, metal, building
materials, clay, etc. or the soil can not be improved by tilling, it must
be improved if trees are to grow. The improvement should consist of removing
some of the existing material and incorporating organic matter such as
compost, peat moss, peat, etc., to improve the soils's permeability. If
the soil is heavy clay, sand should be added by as much as 50% in volume.
If the soil is pure sand, organic loam should be added by as much as 30%
in volume. Gypsum should be added if the soil test indicates high saline
condition also known as a high salt index. Sulfur or aluminum sulfate should
be used to bring down a high pH, although the planting of alkaline tolerant
trees is a better option. Permeability of the soil is essential if water,
air, and roots are to move through the soil.
Tree roots also need warm soil so they can continue to grow all year
round. Even when the soil is frozen, the roots will continue growing until
the soil reaches a temperature of 240F (-5°C). Mulch or forest soil
will protect the roots and usually prevent the temperature from going below
240. Tree tops on the other hand can survive in temperatures as low as
-20°F (-29°C).
Soil Preparation
Roots are opportunistic and will grow in the direction of favorable soil
conditions. These soil conditions include moisture, minerals, oxygen, and
organic matter. How does the municipal arborist provide suitable soil conditions
that will allow the tree's roots to grow and thrive in urban soils?
When transplanting trees into urban soils, arborists should select the
sites containing as much high quality soil as possible. High quality soils
have adequate aeration, moisture, a stable pH, and a good amount of organic
matter. If good soils are not available, amendments such as organic fertilizer
and soil conditioners should be added. Biostimulants, surfactants and water
absorbing gels can be added to the soil. Inocculants of mycorrhizae and
beneficial bacterias as found in products such as Mycor Tree, Tree Saver
Transplant, or equal, are also desirable at planting time. Mature trees
will also benefit from mycorrhizal injections into the soil before, during,
or after a major construction project over their roots.
These tips will encourage tree survival through consideration of root
microorganisms:
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Amend disturbed urban soils with native topsoil when landscaping. Native
topsoil will have mycorrhizal fungi.
-
Maintain a well aerated soil environment because this stimulates mycorrhizal
activity.
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Avoid using vermiculite and perlite as landscape planting amendments because
they inhibit the formation of mycorrhizae essential for tree root development.
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Avoid overusing inorganic fertilizers, especially phosphorus, because they
inhibit the formation of mycorrhizae.
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Avoid unnecessary use of fungicides unless their is a major outbreak of
harmful fungi.
The ideal soil should be 25% air, 25% water, 5% organic matter, and 45%
minerals (soil). By comparison, a typical urban soil is 12% air, 12% water,
1% organic and 75% minerals.
How do you grow trees in downtown soils? How do you compact soil for
sidewalks, curbs, and streets and not compact the soil for street tree
planting? Cornell University has developed a technique called Cornell's
Structural Soil mix. It is basically crushed rock varying in size between
1/2" and 1-1/2" diameter. The rock is easily compressed so any sidewalk
or street construction has the necessary compact soil. At the same time,
the large pores between the rock particles can contain soil, roots, moisture,
and air. The soil is stuck to the rock by using hydrogel as a sticking
agent. The root can grow between the rock particles seeking moisture and
air.
The rock is prepared by being spread out near the construction site.
The hydrogel and soil are added on top of and mixed into the rock. The
ratio is 30 grams of hydrogel to 100 kilograms of soil to 500 kilograms
of rock.
The entire mix is then moved to the construction site so the sidewalk
or street can be built. The trees can be added later. The mix is spread
at least three feet deep and under as much paved surface as feasible. The
mix allows the pavement to be built properly and the trees to grow with
their roots penetrating the voids between the rock particles.
Another growing technique for dealing with poor urban soils is the Amsterdam
Tree Soil which consists of 91% coarse sand, 5% organic matter and 4% clay.
This soil is especially suited for growing trees surrounded by sidewalks.
The sand provides the stable base for the walk construction while tree
roots can utilize the minerals and organic matter.
If the existing soil is so bad that it can not be amended to meet the
ideal conditions, the soil should be completely removed and replaced with
the Cornell Structural Soil mix, the Amsterdam Tree soil, or other soil
improvement techniques. The soil should be amended as necessary before
trees are planted. Attempts to grow healthy trees in poor soil will not
succeed. Where growing conditions are ideal, roots can grow 10 to 15 feet
per year.
Sources:
Bassuk, Nina, "Cornell Structural Soil Mix", City Trees, Vol. 35
No. 1, January/February, 1999. pg. 11. Carlson, Chris R., "Overview of
Urban Tree Root Problems", City Trees, Vol. 31, No. 1, January/February,
1995. P. 19. Chaney, Dr. William, "How Trees Grow", Tree Care Industry,
Vol. VII, No. 4, April, 1996. p. 31. Gillham, Felicia, "Getting to the
Root of Tree Planting", Arbor Age, Vol. 17, No. 11, November, 1997.
p. 8-14. Harris, Richard W., Arboriculture, Prentice-Hall, 1992,
p. 44 48, 150 - 152. Martin, Chris A., "Mycorrhizal Fungi", City Trees,
Vol. 31, No. 4, July/August, 1995. p. 13&14. Marx, Donald H. and Rob
McCartney, "Tree Roots and their Microbial Partners", Arbor Age,
Vol. 17 No. 4, April, 1997. p. 8 - 10. Patterson, Dr. James C., "Soil and
Landscape Design Considerations", City Trees, Vol. 27, No. 1, January/February,
1991. p. 12. Robbins, Wilfred W. et al, Biology, John Wiley &
Sons, 1957. p. 136-153. Watson, Gary W. & Patrick Kelsey, "Soils: The
Root of Tree Problems", Arbor Age, Vol. 13, No. 8, August, 1993.
p. 14 - 18.
Society of Municipal Arborists
Norma Bonham, Executive Director
P.O. Box 11521
St. Louis, MO 63105
Phone: 314-862-3325
Website: http://www.urban-forestry.com/
Trees are the City
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