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Jason R. Montesa 1EPH 1. In previous chapters, growth by apical meristems was described.

In woody species, additional tissues are produced in the stem and root by two other meristems, the _________________ ______________ and the ____________. The new tissues themselves are _____ (_________ ______) and ______________ (______________ and ___________). They are the secondary tissues, and they constitute the plants secondary body. Vascular cambium, cork cambium, wood, secondary xylem, bark, secondary phloem and cork. 2. Name at least three examples of woody plants that are trees and three that are shrubs. Example of woody plants that are trees: sycamores, chestnuts and pines while woody plants that are shrubs are: Roses, Oleanders, and Azaleas. 3. Many herbs live for several years. How do they respond to the bottleneck of the limited conducting capacity of the first-year stem? In some, the first year's leaves die during winter, and during the second year, the plant produces as many leaves as it had during the first year. In other species, adventitious roots are produced that supply conducting capacity directly to the new section of stem being formed. Most of these plants must remain low enough for adventitious roots to reach the soil. These are often rhizomatous, such as irises, bamboo, and ferns. 4. Which groups of plants have secondary growth? Which never do? Gymnosperms and woody angiosperms have secondary growth while herbaceous plants never have secondary growth. 5. Imagine a tree that has a radius of 20 cm and that produces a new layer of wood 0.5 cm thick (the outer radius of the new wood is 20.5 cm and the inner radius is 20 cm). What is the crosssectional area of the new layer of wood (CSA=r2)? If each leaf needs 0.1cm2 of wood to supply it with water, how many leaves can the tree have? If the tree produces a new ring of wood next year that is again 0.5 cm thick, how many leaves can the tree have next year (assume that wood conducts water for only 1 year, not 2 years)? - CSA = x (20.5^2 20.0^2) = 63.6 cm^2 - The tree can accommodate 636 leaves. (63.6cm^2 / 0.1cm^2) - The tree can accommodate 652 leaves (65.2cm^2 / 0.1 cm^2) 6. Woody plants are almost always perennial plants, often living for many years. Describe some advantages of this with regard to a plants ability to occupy a favorable site. Describe some disadvantages with regard to how long a plant must survive pathogens and harsh conditions. An advantage of perennial plants is it tends to occupy their favorable site for a long period of time; moreover it also produces seeds of their own and by their very presence, they prevent the seeds of their competitors from growing at that site. Consequently perennial plants have to

battle insects, fungi and environmental harshness 10, 000 longer and greater. It is also expensive metabolically to construct wood and bark. 7. It must be difficult for secondary growth to arise by evolution. How many times has it evolved in the 420 million years that vascular plants have been in existence? How many of those have become extinct? The secondary growth might have evolved for about 3 times in the last 420 million years and on the present time only 1 of the three have survived. 8. What is the name of the vascular bundles? Between vascular bundles? After the two have formed, what is the shape of the vascular cambium? Is it a series of strips that are aligned up and down the trunk or root, or is it a complete cylinder? Fascicular bundles, interfascicular cambium, complete cylinder 9. Imagine a tree that is growing in springtime. All woody parts that were present last year have a vascular cambium, but all of the twigs that are brand new have just primary growth with vascular bundles. When will a vascular bundle form in these twigs? Will it connect to the old, pre-existing vascular cambium, or will it remain separate? Describe how this occurs. The vascular cambium will form in the twigs when the metaxylem and the metophloem have matured. The vascular cambium will connect to the old, pre-existing vascular cambium. It occurs when the apical meristem extends the axis the new vascular cambium joins at its base with the vascular cambium from the past season. 10. Figure 8-3 is complex but easy to understand. Figure 8-3b shows a tree trunk, and this year is its fourth year of growth. A transverse section at the base of Figure 8-3b would show how many layers (growth rings) of wood (circle 1, 2, 3, 4)? Would a transverse section through the top of Figure 8-3b show any layer of wood this year? Would it show a layer of wood at the end of next year? Would there be any transverse section that does not show the pith and primary xylem; that is, are pith and primary xylem ever lost or destroyed by the formation of wood? The orange layer is labeled Provascular tissue in the top right of Figure 8-3b, but what is it labeled in the middle and lower right diagrams? 4 growth rings, yes, yes, none, it is labeled as the vascular cambium 11. What are the two types of cells in the vascular cambium? Can either type be converted into the other? The two types of cells in the vascular cambium are fusiform initials and ray initials. Yes, fusiform initials can be converted to Ray initials after nuclear division of cells. 12. Look at Figure 8-6. Part (a) shows the lower half of a fusiform initial (actually most fusiform initials would be much taller than this), and part (b) show the same cell after it has divided and one of the daughter cells is developing into a xylem cell. Did the fusiform initial divide with a periclinal wall or an anticlinal wall as it went from part a to part b? Part (d) shows that the same fusiform initial has now produced a second cell that is developing into a phloem cell. Was that

phloem cell produced by a periclinal or an anticlinal division of the fusiform initial? Part (f) show the fusiform initial dividing and forming a second fusiform initial. Is this division occurring by a periclinal wall or an anticlinal wall? The fusiform initial divided with the periclinal wall and the inner cell develops into secondary xylem while the outer cell remains a fusiform initial. The phloem cell was produced by a periclinal division. The division of the fusiform initial and formation of a second fusiform initial was facilitated by the anticlinal wall. 13. Are fusiform initials parenchyma cells, collenchyma cells or sclerenchyma cells? Do they have chloroplast, chromoplasts, or proplastids? Parenchyma, none 14. Look at Figure 8-8. Vascular cambium is not present; it was located far above the top of the micrograph, but you can see there are six rows of wood cells. How many fusiform initials were involved producing visible in this figure? Look at all the cells in any particular row. Were all the cells of that row made by divisions of one fusiform initial or by many different fusiform initials? Six, It was made by one fusiform initial. 15. Are ray initials longer or shorter than fusiform initials? Can they undergo both periclinal and anticlinal divisions or just one or the other? Ray initials are shorter than fusiform initials. Fusiform initials could undergo both periclinal and anticlinal divisions while the ray initials could only undergo the periclinal divisions. 16. Ray initials are typically grouped together in short vertical rows only one cell wide (____), two cells wide (____) or many cells wide (_________). Uniseriate, biseriate, multiseriate 17. True or false: Typically a vascular cambium never has large regions of only fusiform initials or ray initials. True 18. What fraction of cells formed to the interior of the vascular cambium develop into secondary xylem, known as wood- less than half, exactly half, or more than half all cells? More than half all cells develop into secondary xylem. 19. In gymnosperms (conifers like Christmas trees), what types of cells occur in the axial system (the cells produced by fusiform initials)? Which cells are very rare (Hint: Table 8-2)? Wood of gymnosperms is called __________ because a certain type of cell is absent from their axial system. Which type of cell is that? Tracheary elements like tracheids or vessel elements occur in the axial system. Fibers and parenchyma cells are very rare. Softwoods are the wood of gymnosperms. Fibers is absent from their axial system. 20. In basal angiosperms and eudicots, what types of cells occur in the axial system? What is the only type of cel present in the rays of eudicots? The wood of eudicots is called _______ because it has the type of cell lacking in gymnosperms. Which type of cell is that?

Some basal angiosperms mostly have tracheids; however, in the majority of woody angiosperms, all possible types of cells are present. Parenchyma, hardwood, fibers. 21. What types of cells are derived from fusiform initials? What types of cells are derived from ray initials? Is it theoretically possible to have a vascular cambium without ray initials or without fusiform initials? Fusiform initial produces two elongated cells. One continues to be a fusiform initial while the other becomes a cell of secondary xylem or secondary phloem. Ray initials produce short cells, mostly just storage parenchyma and in gymnosperms, albuminous cells. It is not possible to have no fusiform or ray initials. 22. Look at the woods of Figure 8-13. In part (b), all vessels have about the same diameter and are narrow, but the wood in part (c) has some very wide vessels (in the early wood) and some narrow ones (in the late wood, both types produced in the same year). The answer is not in the text, but would you think that these two species have different water conducting needs? Is it possible that one species lives in areas with very rainy spring times and the other lives in an area that is rather dry all the time? Yes, the two species have different water conducting needs, it is supported by the idea that early wood have many large vessels and few fibers while the late wood have abundant fibers and a few narrow vessels. Yes it is also possible that the species live in different areas due to its differences in structure that was caused through its adaptation to the environment. 23. What is a growth ring? How does early wood differ from late wood? Do all species of wood show strong differences between these two phases of a growth ring? Growth rings in secondary xylem are a set wood usually an early wood and a late wood that is produced in one year. The early wood is the wood that is first to be formed it has a high proportion of vessels, while a late wood is formed after the early wood and has a lower proportion of vessels. There are no strong differences between these two phases of growth ring. 24. Why do some people prefer the term growth ring rather than annual ring? What can sometimes happen if a summer is unusually cool? Do you think this occurs more frequently in California or in Alaska? People prefer the term growth ring rather than annual ring because if a summer is usually cool, a tree may fail to grow and produce a ring; these rings are formed but are not truly annual rings. This occurs more frequently in Alaska. 25. If especially wide vessels are produce early in the growing season and only narrow, sparse vessels are produced later, the wood is said to be _______ porous; however, if vessels have similar size and abundance throughout a growth ring, the wood is ______ porous. Name three examples of each type. Diffuse, ring, The examples of diffuse porous are yellow birch, aspen and sugar maple while the examples for ring porous are red oak, sassafras, and honey locust.

26. What changes occur as sapwood is converted to heartwood? The Sapwood will be gradually converted into an inactive wood and it will break its water columns, its parenchyma cells will become resistant to fungi and eventually die. The sapwood is converted to heartwood; moreover a new layer of sapwood is produced annually by the vascular cambium. 27. What is tylosis? How does formation of tyloses slow the spread of fungi in wood? Why doesnt a tree make tyloses in vessels that are still conducting water? Tyloses are plugs that are formed when vessels sops conducting because of cavitation and when adjacent cells push cytoplasm in the vessel through pits. Formation of Tyloses serves to slow the spread of the pathogen through the plant but also restrict water flow, resulting in wilting. In addition, the pathogen causes a general breakdown of the xylem parenchyma cells resulting in various types of gums that add to the blockage. The extent to which fungal toxins are involved in pathogenesis while If a tree make tyloses in vessels that are still conducting water, it may result in killing the tree, because the water contained in the vessels are not permanent and will be used in further processes occurring in the tree. 28. The thickness of sapwood is some indication of how long a growth ring of wood is able to function. Look at Table 8-3. About how long does wood of catalpa function? How about wood of wild black cherry or honey locust? The wood of catalpa is able to fuction for 1 to 2 years while the woods of both wild black cherry and honey locust function for 10-12 years. 29. Because secondary phloem is formed from the vascular cambium just as secondary xylem is, it too has an ________ system and a ________ system. Why do the size, shape, and number of phloem rays match those of xylem rays? Axial, radial, it is a match because both are produced by the same ray initials. 30. In a cross-section of a tree, where are the oldest growth rings-in the outer region or nearer the pith? Where is the oldest secondary phloem-near the outside of the tree or near the cambium? Oldest growth rings are located nearer the pith while the secondary phloem is nearer the vascular cambium. 31. What causes the outermost tissues of a woody stem or root to become pushed outward and expanded? The outward push and expansion of outermost tissues of a woody stem or root are caused by production and differentiation of secondary xylem cells. 32. The layer of cells that produces cork has a technical name and an ordinary name. Give each name. Cork cambiums are produced by the periderm. The cork cambium or phellogen its technical name and bark an ordinary name, divides to produce cells. These cells are then pushed outside and cells with suberin are called cork cells or phellem. 33. Periderm consists of at least two types of cells. Sometimes a third is present. What are these three types of cell? These are cork cambium, layers of cork cells, and phelloderm.

34. Why does a layer or periderm offer only a temporary protection? When the plant makes a new layer of cork cambium, does it make the new layer to the outer side of the failing periderm or to the inner side of it, deeper in the secondary phloem? Its because it is not yet permanent and it can be torn apart, and the surface also should be maintained against the environmental factors that may invade the plant. It makes the new layer to the inner side. 35. In which tissues does the first cork cambium form? When does it usually arise? In which tissues do later cork cambia form? Vascular cambium, it arises before a twig or root is even 1 year old. The first cork cambium may arise in a number of tissues: epidermis,cortex,primary phloem or secondary phloem.Then,later cork cambia form deep in the secondary phloem shortly afterward or in a later time. 36. Are geraniums herbs or woody plants? Do they every form a bark (Hint: Figure 8-23)? Geraniums are woody plants and it is capable of forming a bark. 37. What type of plants produces the cork used for wine bottles and cork board-pines, oaks, maples, eucalyptus, or none of these (Hint: Figure 8-25)? Oak plants are used to produce corks for wine bottles and cork boards. 38. What is the function of lenticels? How are intercellular spaces important for this function? The function of the lenticels is to permit the penetration of Oxygen in all layers of the bark while intercellular spaces are the place where oxygen diffuses. 39. Do roots form wood and bark, or are these secondary tissues present only in stems? Roots form wood and bark, these secondary tissues are also present in roots. 40. Describe the anomalous secondary growth in roots of sweet potatoes. What is the selective advantage of this unusual type of secondary growth? The anomalous secondary growth has more vascular cambia that arise within clusters of vessels in the roots of the sweet potatoes. These cambia acts normally, except the xylem and phloem produced are almost purely parenchyma. The new vessels that were produced may be encircled by another new cambium, and the process is repeated. As the sweet potato becomes quite large, it may contain hundreds of vascular cambia of various age; the secondary tissues are an irregular matrix of parenchyma, a few sieve tubes, some vessels and vascular cambium. The selective advantage is the increase rate of the cell production because instead of having one cambium working to enlarge the root as quickly as possible there are multiples of vascular cambium that works to speed the production of the storage capacity. 41. Describe the formation of included phloem. How did the included phloem of Iresine in Figure 830a become surrounded by xylem?

The formation of included phloem occurs when a secondary phloem is located between two bands of xylem after a new vascular cambium arises in the outermost of the secondary phloem. Furthermore, the Iresine from interior to exterior has an original formation of primary xylem, primary phloem, secondary xylem and secondary phloem. It will raise a new vascular cambium in the outermost part of the secondary phloem and it will produce another formation of primary xylem and phloem, and a secondary xylem and phloem. It will cause the included phloem to be surrounded by the secondary and primary xylem. 42. Describe the secondary growth of monocots like Joshua trees and dragon trees. What are secondary vascular bundles? The secondary growth of monocots occurs when the first two groups undergo a process of anomalous secondary growth. In Joshua trees (some members of the genus Yucca; Fig. 8.32) and dragon trees (in the genus Dracaena), a type of vascular cambium arises just outside the outermost vascular bundles (Fig. 8.33) which originates from cortex cells in the same manner that the interfascicular vascular cambium arises in dicots. This cambium, however, produces only parenchyma; conducting cells are completely absent. Some of the parenchyma cells undergo rapid division and produce columns of narrow cells that differentiate into secondary vascular bundles containing xylem and phloem. The outermost cells of each bundle develop into fibers with thick secondary walls. The parenchyma cells that do not divide like this form a secondary ground tissue, the arrangement of which is almost identical to that of primary tissues. They are "woody" because of the fibers, and they have more conducting capacity and greater strength each year, so branching is feasible. 43. Describe the growth of a palm seedling. For the first few years of life, the seedling becomes wider, but without secondary growth, how is it able to do this? Each adventitious roots adds something. What is it? If the palm stem has 100 vascular bundles and then forms five new adventitious roots with eight bundles each, how many vascular bundles can the stem have above these new adventitious roots? Would the stem be wider or narrower above these adventitious roots? For the first few years of life, the palm trunk becomes wider and the number of leaves increases. It happens without secondary growth. Palm seedlings produce numerous adventitious roots from the base of the short stem in which every root adds extra vascular bundles, and the portion of stem above each new root can have that many more bundles than it does below the root. The palm will have 140 vascular bundles and will be wider. 44. Why can a monocot like an iris branch and increase its number of leaves? Is the fact that the shoot is a rhizome with adventitious roots important? Is water transported from one end of the shoot to the other? Monocots can branch and increase its number of leaves through anomalous secondary growth. Yes, no

45. What is the name of the analysis of tree rings? How is that used to study past climates? How is it used to establish the date when ancient buildings and ships were constructed? Dendrochronology or tree-ring dating is the analysis of tree rings. It is used to study past climates through the reflections of yearly precipitation patterns and those that can be analyzed to reconstruct a record of past climate conditions. Through its growth layers that appear as rings in the cross section of the tree trunk, a record and proof of floods, droughts, insect attacks, lightning strikes, and even earthquakes are seen. The buildings and ships may have wooden beams and its outermost rings may match patterns of the inner rings of living trees and by counting back and finding the age of that set of rings in the living wood, it would also show when the building was established. 46. In tree ring analysis, what is a floating sequence? Why has the sequence for the Middle East been floating, whereas we know the exact dates of each ring in the sequences for North America and Europe? What is so important about the volcanic eruption in 1628 bce? Do you think it is possible that the unusual ring in North American tree sequences was produced by a volcanic eruption in North America and the unusual ring in the Middle East sequence was caused by a different phenomenon that occurred in a different year? If so, then has the floating sequence been anchored? The floating sequence is a tree ring sequence extending over 1503 years and must have begun somewhere earlier than 2200 years BCE. The sequence for the Middle East has been floating because at that time, Middle East was so hot and dry that the cooling caused by the volcanic dust was only enough to produce extra rain and optimal growing conditions rather than frost and poor growing conditions. The volcanic eruption in 1628 BCE produced so much dust that sunlight was blocked around the world, and the summer of 1628 BCE was so cold that frost damage occurred in many trees. Yes, and the sequences have been anchored.

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