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SECONDARY GROWTH
SECONDARY GROWTH IS AN INCREASE IN
GIRTH IN ORGANS WHICH ARE NO LONGER
ELONGATING. THIS INCREASE IN DIAMETER IS
THE RESULT OF ACTIVITY OF TWO CAMBIA:
THE VASCULAR CAMBIUM (VC) AND THE CORK
CAMBIUM (CC). THESE CAMBIA ARE ALSO
PARALLEL TO THE SURFACE OF THE ROOT OR
STEM. THEY FORM A CYLINDER RUNNING THE
LENGTH OF THE STEM OR ROOT. SOMETIMES
THEY ARE CALLED LATERAL MERISTEMS.
SECONDARY GROWTH IS ABSENT IN ALL
SPECIES OF MODERN FERNS AND IN ALL
MONOCOTS (BUT SOME HAVE AN ANOMALOUS
SECONDARY GROWTH) PRESENT IN ALL
GYMNOSPERMS. IT IS PRESENT IN DICOTS
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THAT ARE WOODY AND ABSENT OR OF
LIMITED PRESENCE IN HERBACEOUS SPECIES.
SECONDARY GROWTH OCCURS IN ROOTS AND
STEMS, NEVER IN FRUITS OR FLOWERS. IN
LEAVES THAT ARE EVERGREEN RARELY A
VASCULAR CAMBIUM FORMS, BUT PRODUCES
SMALL AMOUNTS OF SECONDARY PHLOEM
WITHOUT ANY SECONDARY XYLEM IN THE
MIDRIB.
IN SOME MONOCOTS THAT HAVE A TYPE OF
SECONDARY GROWTH A CAMBIUM-LIKE
LAYER PRODUCES MANY SEPARATE
VASCULAR BUNDLES TO THE INSIDE OF THE
STEM AND ONLY PARENCHYMA CELLS ARE
PRODUCED TO THE OUTSIDE.

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VASCULAR CAMBIUM
THE VC OCCURS BETWEEN THE METAXYLEM
AND METAPHLOEM IN VASCULAR BUNDLES. IN
HERBACEOUS SPECIES THIS LAYER
ULTIMATELY STOPS DIVIDING AND
DIFFERENTIATES INTO XYLEM & PHLOEM. IN
WOODY SPECIES THIS DOES NOT HAPPEN AND
THEY CONTINUE TO DIVIDE AND FORM THE
FASCICULAR CAMBIUM. ADDITIONALLY CELLS
BETWEEN VASCULAR BUNDLES RESUME
MITOSIS AND FORM AN INTERFASCICULAR
CAMBIUM. THIS CONNECTS ON EACH SIDE
WITH FASCICULAR CAMBIA. ONCE THIS
HAPPENS THE VC BECOMES A COMPLETE
CYLINDER. THE TERMS FASCICULAR AND
INTERFASCICULAR CAMBIA ARE ONLY USED
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WHEN THE CAMBIA ARE YOUNG AFTER THAT
THEY ARE INDISTINGUISHABLE.
THE VC PRODUCES ONLY TWO TYPES OF
CELLS: FUSIFORM INITIALS AND RAY INITIALS.
FUSIFORM INITIALS ARE LONG TAPERING
CELLS. WHEN IT DIVIDES IT PRODUCES TWO
ELONGATE CELLS. ONE CONTINUES TO BE A
FUSIFORM INITIAL THE OTHER
DIFFERENTIATES INTO SECONDARY XYLEM OR
SECONDARY PHLOEM. IF THE OUTER
DAUGHTER CELL REMAINS AS CAMBIUM, THE
INNER ONE DEVELOPS INTO SECONDARY
XYLEM. IF THE INNER ONE REMAINS AS
CAMBIUM THE OUTER ONE DEVELOPS INTO
SECONDARY PHLOEM. THIS ORIENTATION IS
CONSTANT. WOOD NEVER FORMS TO THE
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EXTERIOR OF THE VC AND BARK NEVER
FORMS ON THE INTERIOR SIDE. IN ANY YEAR
BOTH XYLEM AND PHLOEM ARE PRODUCED
BUT ALMOST ALWAYS MUCH MORE XYLEM
THAN PHLOEM. FUSIFORM INITIALS PRODUCE
THE ELONGATE CELLS OF WOOD TRACHEIDS,
VESSEL ELEMENTS, FIBERS - AND PHLOEM
SIEVE CELLS, SEIVE TUBE MEMBERS,
COMPANION CELLS, FIBERS.
RAY INTIALS ARE SHORT AND CUBOIDAL.
THEY TWO DIVIDE WITH ONE CELL FORMING
XYLEM PARENCHYMA (ON THE INSIDE) AND
ONE CELL REMAINING AS A RAY INITIAL. OR
PHLOEM PARENCHYMA IS FORMED ON THE
OUTSIDE, THE OTHER REMAINING CELL STAYS
A RAY INITIAL. RAY INITIALS PRODUCE SHORT
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CELLS JUST STORAGE PARENCHYMA AND IN
GYMNOSPERMS STRASBURGER CELLS.

SECONDARY XYLEM
ALL CELLS FORMED TO THE INSIDE OF THE VC
DEVELOP INTO SECONDARY XYLEM OR WOOD.
IT IS COMPOSED OF AN AXIAL SYSTEM OF
LONGITUDINALLY ORIENTATED CELLS
DERIVED FROM THE FUSIFORM INITIALS. IT IS
COMPOSED OF TRACHEIDS, VESSELS, FIBERS
AND PARENCHYMA. THIS CARRIES OUT
LONGITUDINAL CONDUCTION OF WATER
THROUGH THE WOOD.
THE RADIAL SYSTEM, DERIVED FROM RAY
INTIALS, CONTAINS ONLY RAY PARENCHYMA
WHICH MAY BE ARRANGED AS UNI-, BI- OR
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MULTI-SERIATE MASSES CALLED RAYS. RAY
PARENCHYMA CELLS STORE CARBOHYDRATES
& NUTRIENTS AND CONDUCT MATERIAL OVER
SHORT DISTANCES RADIALLY WITHIN WOOD.
IN MANY DICOTS THE AXIAL SYSTEM
CONTAINS FIBERS TO GIVE THE WOOD
STRENGTH AND FLEXIBILITY. THOSE WOODS
WITH LARGE AMOUNTS OF FIBERS ARE
CALLED HARDWOODS NOW USED FOR ALL
DICOT WOODS EVEN THOSE THAT LACK
FIBERS AND ARE SOFT (BALSA). WOODS FROM
CONIFERS HAVE FEW OR NO FIBERS AND SO
ARE SOFTWOODS EVEN IF IN SOME CASES
THEY ARE HARDER THAN SOME HARDWOODS.
THE SUCCESSIVE INCREMENTS OF XYLEM
CELLS PRODUCED BY THE VC COMPRISE
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GROWTH LAYERS OR ANNUAL RINGS. THE
CELLS PRODUCED IN THE EARLY PART OF THE
GROWING SEASON (INNER PART) ARE USUALLY
LARGER IN TRANSVERSE SECTIONS THAN
THOSE PRODUCED LATER AND MAY HAVE
THINNER WALLS. THESE CELLS ARE THE
EARLY WOOD. LATE WOOD (=SUMMER WOOD)
PRODUCED LATER IN THE GROWING SEASON IS
COMPOSED OF SMALLER CELLS WITH
THICKER WALLS. AS A RESULT THERE IS
OFTEN A LINE OF DEMARCATION BETWEEN
GROWTH LAYERS.
THE CENTER OF A TREE TRUNK IS USUALLY
DARKER, DRIER AND MORE FRAGRANT. THIS IS
HEARTWOOD. THE LIGHTER, MOISTER OUTER
REGION IS THE SAPWOOD. VESSELS AND
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TRACHEIDS DO NOT CONDUCT WATER
FOREVER. WATER COLUMNS BREAK DUE TO
FREEZING, WIND VIBRATION, TENSION,
INSECTS, ETC. ONCE THE COLUMN BREAKS
TRACHEARY ELEMENTS NEVER CONDUCT
WATER AGAIN. OVER TIME ALL WATER
COLUMNS IN AN ANNUAL RING BREAK. THESE
ARE SEALED OFF WHEN ADJACENT WOOD
PARENCHYMA CELLS PUSH BUBBLE OF
PROTOPLASM THROUGH THE PITS INTO THE
VESSEL FORMING A PLUG A TYLOSE THAT
FILLS THE LUMEN. THE PARENCHYMA CELLS
PRODUCE LARGE AMOUNTS OF PHENOLIC
COMPOUNDS, LIGNIN, WASTE PRODUCTS, ETC.
THAT FILL UP THE LUMEN. THESE INHIBIT THE
GROWTH OF BACTERIA & FUNGI. THE
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HEARTWOOD IS AROMATIC AND DECAY
RESISTANT AND HAS NO LIVING COMPONENTS.
THE YOUNGER OUTER REGION CONTAINS
LIVING PARENCHYMA CELLS AND
FUNCTIONING TRACHEARY ELEMENTS FULL
OF XYLEM SAP HENCE SAPWOOD. A NEW
LAYER OF SAPWOOD IS FORMED EACH YEAR.
ON AVERAGE ONE ANNUAL RING PER YEAR IS
CONVERTED TO HEARTWOOD. THUS
HEARTWOOD BECOMES WIDER WITH AGE;
SAPWOOD HAS A MORE OR LESS CONSTANT
THICKNESS. THIS OCCURS IN OLDER STEMS,
YOUNGER STEMS LACK THESE TWO LAYERS.
WHEN THESE APPEAR DEPEND ON THE
SPECIES: BLACK WALNUT HAS WOOD THAT
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FUNCTIONS FOR TEN YEARS BEFORE
CONVERTING TO HEARTWOOD.

SECONDARY PHLOEM
SECONDARY PHLOEM HAS AN AXIAL AND
RADIAL SYSTEM ALSO. THE AXIAL SYSTEM IS
RESPONSIBLE FOR CONDUCTION UP AN DOWN
THE STEM OR ROOT. IT CONTAINS THE SIEVE
ELEMENTS. FIBER AND NON-CONDUCTING
PHLOEM MAY ALSO BE PRESENT. SIEVE TUBE
MEMBERS & SEIVE CELLS CONDUCT FOR LESS
THAN A YEAR WITH ONLY THE INNER MOST
LAYER OF PHLOEM CAPABLE OF CONDUCTION.
THE RADIAL SYSTEM IS COMPOSED OF
PHLOEM RAYS MADE UP OF RAY PARENCHYMA
CELLS THAT ARE USED FOR STORAGE.
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AS THE VC CREATES NEW XYLEM & PHLOEM
THE PRESSURE IT PUTS ON THE DELICATE
CELLS OF THE PRIMARY PHLOEM
ESSENTIALLY CRUSH THEM OUT OF
EXISTENCE. ONLY THE THICK WALLED
PRIMARY PHLOEM FIBERS REMAIN INTACT.
THE CONTINUED EXPANSION OF THE VC AND
XYLEM RESULTS IN THE OUTWARD
DISPLACEMENT OF THE PRIMARY PHLOEM
AND SECONDARY PHLOEM. THESE ARE
COMPRESSED AND CRUSHED. SO ONLY THE
INNER 1-2 LAYERS OF PHLOEM REMAIN
FUNCTIONAL.
CAMBIAL ACTIVITIY HS A DEEP EFFECT ON
THE LIVING TISSUE TO THE EXTERIOR OF THE
SECONDARY XYLEM; IT HAS LITTLE OR NO
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EFFECT ON THOSE TISSUES TO THE INTERIOR.
THE PRIMARY XYLEM, PITH, AND LAYERS OF
SECONDARY XYLEM REMAIN MORE OR LESS
INTACT. THESE NON-LIVING CELLS WITH
THICK WALLS RESIST THE FORCES EXSERTED
AGAINST IT.

CORK CAMBIUM
THE PRODUCTION OF SECONDARY XYLEM
CAUSES THE VC AND SECONDARY PHLOEM TO
BE PUSHED OUTWARD. PRODUCTION OF NEW
PHLOEM ALSO CONTRIBUTES TO THE
INCREASED DIAMETER OF THE ROOT OR STEM.
TISSUES ON THE PERIPHERY OF THE PLANT
EITHER NEED TO GROW IN CIRCUMFERENCE
OR BE TORN APART. BOTH DO HAPPEN BUT
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BOTH MUST BE CONTROLLED. THE INTEGRITY
OF THE PLANTS SURFACE MUST BE
MAINTAINED TO PROTECT AGAINST WATER
LOSS AND PATHOGEN INVASION.
RAY STORAGE PARENCHYMA CELLS OF THE
SECONDARY PHLOEM BECOME RE-ACTIVATED
AND UNDERGO DIVISIONS RESULTING IN A
NEW CAMBIUM THE CORK CAMBIUM (=
PHELLOGEN).
THE CORK CAMBIUM HAS CELLS THAT ARE
ALL CUBOIDAL. AFTER EACH DIVISION THE
INNER CELL ALWAYS REMAINS CORK
CAMBIUM. THE OUTER CELL BECOMES A CORK
CELL (= PHELLEM CELL). IN SOME PLANTS THE
CORK CAMBIUM MAY PRODUCE A 1-2 CELL
LAYER TO THE INSIDE WHICH MATURES INTO
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A LAYER OF PARENCHYMA CALLED THE
PHELLODERM.
THE CORK CAMBIUM + LAYERS OF CORK
CELLS + PHELLODERM (IF ANY) = PERIDERM.
MATURING CORK CELLS HAVE PRIMARY CELL
WALLS ENCRUSTED WITH SUBERIN MAKING
THEM WATER PROOF; THEN THEY DIE. THE
PERIDERM IS SO IMPERMEABLE THAT ALL
PLANT MATERIAL EXTERNAL TO IT
(EPIDERMIS, CORTEX, OLDER SECONDARY
PHLOEM) DIES FOR LACK OF WATER.
PERIDERM OFFERS ONLY TEMPORARY
PROTECTION BECAUSE THE ROOT/SHOOT
CONTINUES TO GROW IN GIRTH STRETCHING
THE PERIDERM. CORK CAMBIUM IS USUALLY
SHORT-LIVED, LASTING FOR A FEW WEEKS. IT
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THEN BECOMES CORK AND DIES. LATER A NEW
CORK CAMBIUM FORMS IN YOUNGER
SECONDARY PHLOEM AND THE CYCLE STARTS
OVER. OVER TIME MANY LAYERS OF CORK
CAN BE BUILT UP.
THE OUTER BARK IS ALL TISSUES OUTSIDE
THE INNERMOST CORK CAMBIUM. ALL
SECONDARY PHLOEM BETWEEN THE VC AND
THE INNERMOST CORK CAMBIUM IS THE
INNER BARK. THE AMOUNT OF BARK FORMED
IS QUITE VARIABLE. IT IS CONTINUOUSLY
FALLING OFF THE TREE.

ROOTS
LATERAL ROOT DEVELOPMENT IS
ENDOGENOUS ARISING FROM DEEP SEATED
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TISSUE. IN SEED PLANTS THIS IS THE
PERICYCLE A REGION BETWEEN THE
PRIMARY VASCULAR TISSUES AND THE
ENDODERMIS. IN THE FERNS LATERAL ROOTS
DEVELOP FROM THE ENDODERMIS & CORTEX.
IN SEED PLANTS LATERAL ROOT PRIMORDIA
ARE INITIATED IN THE PERICYCLE PROXIMAL
TO THE ZONE OF ELONGATION. THIS IS
APPARENTLY UNDER THE CONTROL OF
HORMONES (AUXIN, CYTOKININS, ETHYLENE)
AND GENES. CELL DIVISIONS IN THE
ENDODERMIS RESULT IN THE FORMATION OF
A SHEATH OVER THE DEVELOPING LATERAL
ROOT AS IT PUSHES ITS WAY THROUGH THE
CORTEX. A ROOT CAP, PROCAMBIUM, GROUND
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MERISTEM AND PROTODERM DIFFERENTIATE
IN THE APICAL REGION.
CONNECTIONS BETWEEN LATERAL ROOTS
AND PARENT ROOTS DEVELOP BY
DIFFERENTIATION OF VASCULAR TISSUES
WITHIN PARENCHYMA TISSUES IN THE
INTERVENING REGION BASICALLY THE
FORMATION OF A VASCULAR PLEXUS. THIS IS
A MIXTURE OF MANY SMALL, SHORT
TRACHEARY ELEMENTS CONNECTED BY
BORDERED PITS. THIS PROVIDES AN
EFFECTIVE APOPLASTIC CONDUIT FOR WATER
& SOLUTES.
ADVENTITIOUS ROOTS MAY ORIGINATE FROM
CALLUS TISSUE ASSOCIATED WITH WOUNDS,
OR ENDOGENOUSLY FROM PERICYCLIC
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PARENCHYMA, FROM INTERFASCICULAR
PARENCHYMA OR AXIAL PARENCHYMA IN
SECONDARY PHLOEM.

ROOTS & SECONDARY GROWTH
ROOTS OF GYMNOSPERMS AND WOODY
DICOTS UNDERGO SECONDARY GROWTH, AS
DO THE STEMS. A VC ARISES IN THE SAME WAY
AS IT DOES IN THE STEM. THE NEW VC HAS THE
SAME STAR SHAPE AS THE PRIMARY XYLEM
BUT IT SOON BECOMES ROUND AS THE
CAMBIUM IN THE SINUSES OF THE PRIMARY
XYLEM PRODUCES MORE SECONDARY XYLEM
THAN REGIONS OF VC NEAR THE ARMS.
EVENTUALLY A CIRCULAR VC IS PRODUCED
AND UNEQUAL GROWTH STOPS.
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WOOD IN MANY ROOTS IS VERY MUCH LIKE
THAT OF THE STEM. PERENNIAL ROOTS ALSO
FORM BARK. THE CORK CAMBIUM USUALLY
ARISES IN THE PERICYCLE CAUSING THE
ENDODERMIS, CORTEX AND EPIDERMIS TO BE
SHED. THE PRIMARY PHLOEM CAN BE
COMPRESSED AND OBLITERATED.
IN HERBACEOUS PERENNIALS SECONDARY
XYLEM MAY BE PRODUCED IN LIMITED
AMOUNTS, BUT NO OR LITTLE SECONDARY
PHLOEM IS PRODUCED. THE METAPHLOEM
FUNCTIONS AS THE PIPELINE FOR
PHOTOSYNTHATES THROUGHOUT THE LIFE OF
THE PLANT. IN THOSE PLANTS A PERIDERM
DOES NOT DEVELOP AND THE CORTEX IS
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RETAINED. THE EPIDERMIS DEVELOPS A
THICK CUTICLE AND AN EXODERMIS FORMS.
IN ROOTS WITH LARGE AMOUNTS OF
SECONDAY VASCULAR TISSUE A CORK
CAMBIUM DIFFERENTIATES FROM THE
PERICYCLE. IT PRODUCES PHELLEM TO
EXTERIOR AND PHELLODERM TO THE
INTERIOR RESULTING IN A PERIDERM THAT IS
CONTACT WITH THE SECONDARY PHLOEM.
THE CUT OFF CORTEX AND EPIDERMIS DIE
AND SLOUGH OFF. A PERIDERM LIKE THAT OF
A STEM DEVELOPS. HOWEVER, IN SOIL THE
OUTER PERIDERM DECAYS SO THE PERIDERM
REMAINS RELATIVELY THIN AND THE ROOT
SURFACE SMOOTH.