BIO203 Lecture 6: Secondary Growth & Nitrogen

As plants increase in height why don’t they fall over?

- Secondary growth is all about the fact that as plants get taller, they do not fall over
because they get wider as well
- They are wider at the bottom than they are at the top which is a great way of supporting
- They also have an anchoring system (the roots)

Plants increase in girth & undergo structural reinforcement

- Not only are they getting wider, that trunk is becoming woody
- Xylem is not only strong enough to resist the implosion from negative pressure, its
basically a bunch of strongly reinforced tubes that can support weight
- So, strengthening and increase in width, all done by secondary growth
o Plants compensate for increase in size by adding bulk and reinforcement to their
stems
- Primary growth is from the shoot apex and the root apex
o From apical meristems
- Secondary growth is getting wider (increase in girth) it is due to meristem called The
Cambium
o There are two types:
 1 ) vascular cambium which makes the plumbing (phloem on outside and
xylem on the other)
 2) the cork cambium which makes the protective bark on the outer layer
and they are around the circumference of the trunk
- Secondary growth can grow not just in trees but also herbaceous species towards the
end of the growing season
o Like in tomato plants if you cut it, you will see that it is woody in there
- Monocots do not do secondary growth

Thick walled cells within xylem constitute major proportion of structural reinforcement
- These xylem vessels and tracheid are the reinforcement
o Strong cell types, sclerenchyma with secondary walls composed of lignin:
tracheid, fibres, vessel elements i.e. elements of xylem
o They are wood
- Most of the diameter of the tree is xylem
o With some reinforcement of some fibers and things and lignin

Increasing the diameter of the plant

- Increase in girth occurs predominantly by the addition of new xylem to the primary stem
to make a secondarily-thickened stem
o The new xylem is laid down around the whole circumference of the plant
o The new xylem needs to be made, so the meristems divide and then those cells
differentiate into what they are supposed to be
  On the outside they are becoming phloem and on the inside they are
becoming xylem
- Most of this growth is lateral/periclinal division
o Increasing the girth by generating cells inside and outside that meristem

Secondary growth
- Under the microscope picture, there is still discrete. (look at the pic)
- Fascicular cambium (pro-cambium): making new cells within vascular
- You start to get secondary growth when the parenchyma became a meristematic
o Interfascicular cambium is what they are called
o When that is done forming you have the vesicular cambium
 Together they form a continuous ring around the stem
 Called the cambial ring

The Cambial Ring

- There is this transition from strands to continuous that makes xylem and phloem
throughout the entire circumference of the plant
o Before discrete strands

Radial growth
- The first division makes another potential meristematic cell
o To maintain the population
- Then one of them can differentiate and become a xylem cell
- Then another division of the meristem to make 2
o That one will become a phloem cell
- So, you have division that is maintaining the population of meristems
o The division generated daughter cells that can become something
o Depending on which side of the meristem it is, you can get them phloem on the
outside or xylem on the inside
- This is a periclinal division (longitudinal)
o If you just have those, there is a problem
 You are generating cells on the inside which means you are increasing the
diameter, and therefore the circumference and you will need to increase
the number of meristems to go all around
 Every now and again, you need to make cells between cells of a meristem
so that the circumference keeps pace with the increasing circumference
as you are generating cells
 This is an anticlinal division
o Most of it is periclinal, but there has to be some anticlinal or you would end up
with canyons appearing on the tree
Cambium produces secondary xylem (wood) to inside of this layer increasing girth
- Three year tree
- The very thin line is the cambium
- Dicot or gymnosperm?
o If you look at the wood of a dicot you will see vessels and tracheid
o If everything looks the same except for a few holes, it is a gymnosperm
o So, this one is most likely and angiosperm
o Only this year’s phloem is working, so the 3rd, the rest is dead inner bark
 It is dead and filled with metabolites that stop microbes from coming
through
o Xylem could all be functioning
 The most recent is the outermost

Gymnosperm wood
- They are uniform expect for these holes which are called resin ducts
o The function of them is to protect from damage and microbes getting in

From primary to secondary growth

- Right under the apical meristem
o You got some area that will be vascular strands, right now they are not
differentiating
o The cells in there are a bit more elongated than their neighbours and they will
elongate further and become phloem and xylem
- More cells are made by that meristem and they will continue to grow and become
something and then they might be completely differentiated and then they became
oriented into that secondary growth arrangement
- The ones further and further away are undergoing different types of differentiation
o Eventually you will have proper phloem and xylem that is being formed
 At this point you might still have some growth, so you get protoxylem
o Eventually you will get that interfascicular cambium
o And then you will get the secondary growth
 After one year of growth you will start to get something that looks like
twig or branch with woody part on the outside
 Every part of part of the tree is doing this
o And then you have the cork cambium coming in and it is basically the bark

Why are there annual rings?

- In the tropical trees you do not get it
o Unless there is a discrete dry and wet seasons they are no annual ring
- They are usually about the annual/seasonal changes between the availability of
resources and conditions that enable a plant to grow
 o Ex. Spring, summer you get a lot of growth and a high-water demand and a lot of
xylem being formed. And then you have winter and there is dormancy, no cells
being made by that meristem but as it goes to stopping you have less and fewer
cells being made. sot you have a ring of smaller ring
- Dendrochronology: Dating based on tree rings.
o We can line up the width and the growth that happens in those rings
o Also climate change; you can see what has happened
- The way those rings are laid down:
o What the rings are made of, tracheid, xylem vessels, how much lignin is in them
o Look at slide 23
o The way that the wood is laid down has a big impact on the structural qualities of
the wood and how humans use it

Heartwood and sapwood

- Labeling diagram on the exam
- Where are you most likely to find inactive old xylem? The inside/heartwood

Secondary growth in 3D
- The direction in which water is being transported
- The mature xylem and phloem still need a source for water, nitrogen, and phosphorus
- It is transported to them through the ray cells
o They are taking water and nutrients out of the xylem and making sure that it is
made available to those cells that need it
- When you make new xylem and phloem you have to make new transport system
o So you need to make new ray cells
- Two cell types in vascular cambium:
o Fusiform initials - give rise to elements of secondary xylem and phloem - the
axial transport system (the plumbing)
o Ray initials - give rise to rays (ray cells)- the radial transport system
 Multiseriate ray (several cells wide)
 Uniseriate (one cell wide)

Rays: Living parenchyma cells

- They move things out of the xylem to the living portion of the trunk
- They are also the place where nutrients might be stored for the winter or the dry season
and remobilized when the cambium needs to start growing

Bark formation (the cork cambium)

Secondary phloem (inner bark)

- Phloem is much thinner in cross-section, only functional for about a year and the older
parts of it tend to get crushed and become small bits of cell debris
- Much less of it than secondary xylem
 - Typically contains many fibers
- Youngest cells are closest to the cambium, oldest become crushed
- Only most recently formed cells are active

Cork (outer bark)

- Is so variable that it depends on the species, so they will differ in structure: the depth,
structure etc.
- It depends on the cork cambium and how it is generating the cells, how they are
differentiating and any differential rate in that as the circumference of the plant
increases. Also, what secondary metabolites, what lignin, waxes, are being put into
those cells before they die and become protective layer
- What is happening is new cells are being generated, growing, they are differentiating,
filled with secondary metabolites that basically waterproof and kill microbes and then
they die

Cork formation
- The cork cambium forms underneath the epidermis
o The meristematic layer doesn’t form on the outside of the stem
- What happens to the epidermis? It is toast and it will die. There is a cuticle layer on top
o It will then get isolated from any source of nutrient or sugars b/c underneath it
that cork cambium is generating new cells whose destiny is to become dead and
waterproof and no nutrients will be able to make it through to the epidermis
o Epidermis is just there to protect the plant stem while it is young but only until
secondary growth happens and bark forms
- The cork cambium is on the outside of the inner bark (inner bark is generated by the
dead phloem so basically the vascular cambium)
- So the order : xylem, vascular cambium, phloem, dead phloem, cork cambium, cork
bark)
*do not remember phelloderm*
- So for the cork cambium, it is like the vascular in that you have regular cells that all of a
sudden become meristematic
o They are making cells in one direction, just on the outside, unidirectional
o The cells fill up with a bunch of metabolites and stuff
o As the plant is growth its girth is increasing so the cork cambium is not covering
it anymore so, a whole new cork cambium is often initiated below existing cork
layers to compensate for increasing width
 It is the reason why you have different structure barks because you are
pushing those old bark layers further outside and as the circumference
increases, that bark will break apart and become non-continuous
 If you add up those broken piece, it will equal to the circumference of the
plant when it was younger
 Why you get crevasses – you are making a greater circumference now
than then, but it is still there
Lenticel cells
- They form above where there was a stomatal port on the stem when it was green and
they maintain their gas permeability that bacteria will stck to but not get into but gases
can still be exchanged
- The cork contains waxy substances (e.g. suberin) which make the bark impermeable to
water and gases.
- To allow the outer cells to metabolize (they need oxygen) there are gaps in the cork

Monocots do not have secondary growth

- Outside of the trunk of the tree is leaf scars
- There is some cell division after it sheds the leaf and lignification and that is what
increases the width
- But no discrete meristem doing anything
o They do not live as long b/c they cannot make new cells forever
o Pith fills up with new vascular cells every time the monocot makes a new root
and the shoot makes the connection all the way down
o It also does not shut down its phloem every year like the dicot, it is viable for a
few more years and that helps it live long enough