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For review, what is the function of the xylem?
the tissue that transports water and minerals from the root system to the aerial portions of the plant
For review, what is the function of the phloem?
the tissue that translocates the products of photosynthesis -- sugars -- from mature leaves to areas of growth and storage, including the roots
The cells of the phloem that conduct sugars and other organic materials throughout the plant are called..
sieve elements
Sieve element is a comprehensive term that includes both the highly differentiated ___________ of the __________ and the relatively unspecialized __________ of ___________
sieve tube elements; angiosperm; sieve cells; gymnosperms
The phloem tissues contains what kind of cells?
companion cells and parenchyma cells
________ is translocated in the phloem sieve tube elements
sugar
Mature sieve elements are living cells specialized for...
translocation
What are some characteristics of sieve tube elements? (i.e what are they lacking)
- they lose their nuclei and tonoplast during development
- microfilaments, microtubules golgi bodies, and ribosomes are generally absent from mature cells
- modified mitochondria, plastids, and smooth ER
- nonlignified walls
How are damaged sieve elements sealed off?
- short term mechanisms involve sap proteins
- long-term mechanisms involve closing sieve plate pores with callose, a glucose polymer
The main phloem proteins involved in sealing damaged sieve elements are structural proteins called..
P-proteins
- absent in gymnosperms
What is a longer-term solution to sieve tube damage?
production of the glucose polymer, callose
Each sieve tube element is usually associated with one or more _________
companion cells
What role do companion cells play?
they play a role in the transport of photosynthetic products from producing cells in mature leaves to sieve elements in the minor veins of the leaf
- take over some of the critical metabolic functions, such as protein synthesis, that are reduced or lost during differentiation of the sieve elements
What are the 3 types of companion cells?
ordinary companion cells
transfer cells
intermediary cells
all have dense cytoplasm and abundant mitochondria
___________ have chloroplasts with well-developed thylakoids and a cell wall with a smooth inner surface
ordinary companion cells
_________ are similar to ordinary companion cells, except for the development of fingerlike wall ingrowths, particularly on the cell walls that face away from source sieve elements
transfer cells
____________ appear to be well suited for taking up solutes via cytoplasmic connections.
intermediary cells
What is the most characteristic feature of intermediary cells?
the presence of many plasmodesmatal connections to surrounding cells, having many small vacuoles, and poorly developed thylakoids
Where are transfer cells found in plants?
where transport sugars enter the apoplast during the movement of sugars from mesophyll cells to sieve tube elements
What do transfer cells do?
transport sugars from the apoplast to the symplast of the sieve elements and companion cells in the source
What do intermediary cells do?
they function in symplastic transport of sugars from mesophyll cells to sieve elements
What is the function of ordinary companion cells?
they can function in either symplastic or apoplastic short-distance transport in source leaves, depending in part on plasmodesmatal frequencies
Sap is translocated from area of supply called __________ to areas of metabolism or storage called __________
sources; sinks
What do sources include?
exporting organs, typically mature leaves that are capable of producing photosynthate in excess of their own needs
- photosynthate refers to products of photosynthesis
What do sinks include?
all nonphotosynthetic organs of the plant and organs that do not produce enough photosynthetic products to support their own growth or storage needs.
ex: roots, tubers, developing fruits, and immature leaves
What is the pattern of transport in the phloem?
source-to-sink movement
What is the most abundant substance in the phloem?
water
What is the dissolved in the water in the phloem?
translocated solutes -- carbohydrates, amino acids, hormones, some inorganic ions, RNAs, proteins, secondary compounds, etc
What is the most common sugar transported in the sieve elements?
sucrose
the rate of movement of materials in the sieve elements can be expressed in two ways:
as velocity or as mass transfer rate
The most widely accepted mechanism of phloem transport in angiosperms is...
the pressure-flow model
What is the pressure flow mode?
It explains phloem translocation as a flow of solution driven by an osmotically generated pressure gradient between source and sink
The movement of photosynthate (products of photosynthesis) into sieve elements is called...
phloem loading
The movement of photosynthate (products of photosynthesis) from sieve elements to sink cells is called...
phloem unloading
What establishes the pressure gradient of the pressure-flow model?
phloem loading at the source and phloem unloading at the sink
What are the 3 different mechanisms that exist to generate high concentrations of sugars in the sieve elements of the source?
- photosynthetic metabolism in the mesophyll
- conversion of photoassimilate to transport sugars in intermediary cells (polymer trapping)
- active membrane transport
In source tissues, an accumulation of sugars in the sieve elements generates a...
low negative solute potential and causes a steep drop in water potential
According to the pressure-flow model, movement in the translocation pathway is driven by..
transport of solutes and water into source sieve elements and out of sink sieve elements
There is no ________________ in single sieve tube elements, and solutes and water move at the _________ velocity
bidirectional transport; same
The energy requirement for transport through the phloem pathway is __________ in herbaceous plants
small
In the process called ______________, sugars are transported into the sieve elements and companion cells
phloem loading
In the phloem loading, once inside the sieve elements, sucrose and other solutes are translocated away from the source, a process known as..
export
Translocation through the vascular system to the sink is referred to as..
long-distance transport
Sucrose moving from producing cells in the mesophyll to cells in the vicinity of the sieve elements in the smallest veins of the leaf is referred to as..
short-distance transport
Phloem loading can occur via the..
apoplast or symplast
What is the process for apoplastic loading?
The sugars enter the apoplast near the sieve element - companion cell complex. Sugars are then actively transported from the apoplast into the sieve elements and companion cells by an energy-driven, selective transport located in the plasma membranes of these cells.
Apoplastic phloem loading leads to three basic predictions:
1. transported sugars should be found in the apoplast
2. in experiments in which sugars are supplied to the apoplast, the exogenously supplied sugars should accumulate in sieve elements and companion cells
3. inhibition of sugar efflux from the phloem parenchyma or of uptake from the apoplast should result in inhibition of export from the leaf
Sucrose uptake in the apoplastic pathway requires...
metabolic energy
The fact that sucrose is at a higher concentration in the sieve element-companion cell complex than in surrounding cells indicates that...
sucrose is actively transported against its chemical-potential gradient
Phloem loading in the apoplastic pathway involves a..
sucrose-H+ symporter
A __________ pathway has become evident in species that transport raffinose and stachyose, in addition to sucrose
symplastic
What is the polymer-trapping model?
the model states that the sucrose synthesized in the mesophyll diffuses from the bundle sheath cells into the intermediary cells through the abundant plasmodesmata that connect the two cell types
In the intermediary cells, raffinose is sythesized from sucrose and galactinol, thus maintaining the diffusion gradient for sucrose. Because of its larger size, raffinose is not able to diffuse back into the mesophyll
Raffinose is able to diffuse into the sieve elements. As a result, the concentration of transport sugar rises in the intermediary cells and the sieve elements.
The polymer-trapping model makes three predictions:
1. Sucrose should be more concentrated in the mesophyll than in the intermediary cells
2. The enzymes for raffinose and stachyose synthesis should be preferentially located in the intermediary cells
3. The plasmodesmata linking the bundle sheath cells and the intermediary cells should exclude molecules larger than sucrose. Plasmodesmata between the intermediary cells and sieve elements must be wider to allow passage of raffinose and stachyose
What are the steps that are involved in the import of sugars into sink cells?
1. phloem unloading
2. short-distance transport
3. storage and metabolism
In ______ organs, sugars move from the sieve elements to the cells that store or metabolize them
sink
A transition from sink to source status occurs later in development, when the leaf is approximately ______% expanded, and it is usually completed when the leaf is ______ to ______% expanded.
25; 40 to 50
What does Figure 11.19 represent?
Autoradiographs of a leaf of summer squash.
A) The entire leaf is a sink, importing sugar from the source leaf
B-D) The base is still a sink. As the tip of the leaf loses the ability to unload and stops importing sugar (as shown by the loss of black accumulations) it gains the ability to load and to export sugar
What is an apoplast"?
collective cell wall sapce
What is symplast?
the collective cytoplasm
A ___________ pathway has become evident in species that transport _______ and _________
symplastic; raffinose; stachyose
The driving force for long-distance PHLOEM transport is
Pressure created by accumulated solutes and osmosis
Briefly explain apoplastic phloem loading (loading from the cell-wall space; pictures rarely hurt)
a. phloem loading from the cell wall space
b. sucrose - H+ symporters use PMF to accumulate sucrose into the companion cells
c. accumulating sucrose in the companion cell and needs energy
d. sucrose goes against its concentration gradient, and H+ goes down the concentration gradient
Briefly explain symplastic phloem loading
- sucrose cannot escape
- plasmodesmata connect mesophyll all the way to the sieve elements
- accumulation of sugar is by converting it into larger saccharides that can't diffuse back through the plasmodesmata
- does not require ATPase
What is true of phloem sieve tubes
They are membrane lined
Which of the following is common to both cellular respiration in mitochondria and the light reactions of photosynthesis in plastids?
oxygen is one of the byproducts
Classify the following as either a source tissue or a sink tissue (use the words "source" or "sink only, without quotation marks).
Cotyledon in a developing seed
Cotyledon in a germinating seed
Mature leaf
Immature leaf
Mature flower
Immature flower
- sink
- source
- source
- sink
- source
- sink
A tomato fruit growing on a vine will receive most of its nutrients and water from
The mature leaves closest to it
What is true of phloem companion cells
- They are cytoplasmically dense
- they are closely associated with sieve elements
If you analyzed the phloem sap of a plant and found high concentrations of ________________, the plant most likely phloem loads through the symplasm
raffinose family oligosaccharides
If I had a compound that specifically inhibited the activity of plasma membrane ATPases, and if I observed that this compound resulted in an immediate cessation of phloem transport when applied to a plant, what type phloem loading is most likely being employed?
apoplastic phloem loading
Phloem transport of sucrose is expected to remain fairly constant during diurnal cycles (day/night cycles) because
Sucrose and starch are produced during the day, and starch is converted to sucrose at night
What is the driving force for phloem transport?
Pressure created by accumlated solutes and osmosis
What are examples of source?
- cotyledon in germinating seed
- mature leaf
- mature flower
What are examples of sink?
- cotyledon in developing seed
- immature leaf
- immature flower
Fruit and young leaves get nutrients from the ________ and go in all directioins
phloem