Understanding Cambial Behaviour. The key to wood quality
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1 Understanding Cambial Behaviour The key to wood quality
2 A brief history Terminology Dormancy and reactivation Growth of derivatives and wall formation Pitting and plasmodesmata
3 A brief history
4 Nehemiah Grew ( )
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7 Grew s drawing of elm (detail)
8 Charles Francois Brisseau-Mirbel Proposed that cambium was a tissue rather than a sap (1808)
9 Mirbel s (1827) diagram of elm (from Larson, 1994)
10 Cambial cell theories Hartig (1853)- Back to back theory Phloem initial Xylem initial
11 Cambial cell theories Sanio (1863)- Single initial theory
12 Cambial cell theories Raatz/Mischke (1892) Multiple initial theory
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14 Oblique orientation of plane of division Kinoplasmic fibres (microtubules) Kinoplasmosomes (phragmoplast
15 From Bailey (1923) Anticlinal pseudotransverse division Transverse division
16 Length of cambium/cambial age (from Bailey 1923) A: Conifer or vessel-less dicot. (12 species) B: Less specialised dicot. (10 species) C: Highly specialised dicot. (10 species) D: Dicot. with storeyed cambium (10 species)
17 Vacuolation in cambial cells (Bailey 1930)
18 Pinus radiata Active cambium Nomenclature Cambium? Cambial Zone?
19 Butterfield (1975) IAWA Bulletin Cambium a multiseriate zone of periclinally dividing cells lying between the differentiating secondary xylem and phloem, with a distinct initial capable of both periclinal and anticlinal divisions lying somewhere within each radial file of cells
20 Cambium according to Butterfield
21 Schmid (1976) IAWA Bulletin Cambium equivalent to the initiating layer Cambium applied to the entire differentiating region might lead to the conception that the cambium is a multiseriate layer of initials
22 Cambium according to Schmid?
23 The difficulty of identifying the initial means the terms have been used interchangeably
24 Pinus radiata Mid-winter A slowly dividing meristem
25 Pinus radiata Mid-winter Cambium Butterfield Schmid?
26 Humpty Dumpty From Through the Looking Glass and what Alice found there by Lewis Carroll When I use a word, Humpty Dumpty said in rather a scornful tone, it means just what I choose it to mean neither more nor less
27 Can an initial ever be identified with certainty?
28 Aesculus hippocastanum February Cells appear similar across the cambium
29 Identifying an initial Boundary parenchyma phloem side Boundary parenchyma xylem side
30 A. hippocastanum TEM Boundary parenchyma (phloem side)
31 A. hippocastanum TEM Phloem cells in suspended or slow development
32 A. hippocastanum TEM Fusiform Initial
33 A. hippocastanum TEM Boundary parenchyma xylem side
34 Sieve element/companion cell pair in a state of arrested development Initial
35 Boundary parenchyma Companion cell precursor Sieve/element precursor
36 Phloem boundary cell Previous season s phloem: Sieve tube member Companion cells 9 February
37 Dormancy and reactivation
38 Dormant Cambium Fragmented vacuome
39 Storage materials in dormant fusiform cells Spherosomes (lipids) Protein bodies Thick cell walls
40 9 February Fusiform initial Ray initial Starch
41 Cambial reactivation in Aesculus Activity can be detected in the cytoplasm of cambial zone cells long before any signs of activity are displayed by the tree.
42 Active dictyosome in a boundary layer cell of dormant cambium 23 February
43 Developing and mature coated vesicles 8 March
44 Reactivation (16 March) Expanding phloem precursors Fusiform initial Boundary parenchyma
45 Dividing phloem mother cell (16 March) New tangential wall
46 13 April Boundary parenchyma Cytoplasm confined to a thin parietal layer Boundary parenchyma
47 23 April Developing phloem cells Dividing initial Xylem mother cell New xylem elements
48 Typically in the Reading area, Aesculus budbreak occurs in late March, with leaves fully emerged by late April Xylem formation appears to begin coincidentally with leaves beginning to export photosynthate
49 Reactivation sequence Larson (1994): Xylem production first 26 species Phloem production first 21 species Simultaneous production 10 species
50 Observations are inconsistent between authors Acer pseudoplatanus, Quercus rubra, Pinus sylvestris and Vitis vinifera appear in the list of xylem reactivators and phloem reactivators In the Pinaceae: Pinus halepensis and rigida are xylem reactivators Pinus banksiana, resinosa, and strobus (five authors) are phloem reactivators Picea excelsa, rubens and rubra are xylem reactivators, Picea abies is a phloem reactivator Picea glauca a simultaneous reactivator.
51 Phloem production in Aesculus Phloem annual growth rings marked by boundary parenchyma The number of phloem cells in each file is similar to the number of overwintering precursors All the phloem for the season is produced at the beginning of the season
52 Pinus radiata Active cambium producing both xylem and phloem throughout the season
53 Growth of derivatives and wall formation
54 Cell enlargement Quercus robur
55 20 April Developing xylem cells Boundary parenchyma Previous year s latewood fibre
56 Cell tip growing between fibres
57 Enlarging vessel element Boundary parenchyma
58 Developing fibres are compressed and files of cells distorted by vessel enlargement
59 Perforation plates
60
61 Secondary wall formation This the classic representation of the wall of a cell that has all possible wall layers: the ML+P+S1+S2+S3 +HT+W wall zones
62 Cell wall Layers Middle lamella Primary wall S1 S2 S3 Helical thickening (tertiary layer) Cell lumen
63 Earliest stages of cellulose deposition forming the S1 layer
64
65 Microtubules and cellulose orientation
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69 Tubulin in a fusiform cambial cell of Aesculus (B), and developing fibres (C, D, E)
70 Tubulin in developing fibres in Populus
71 Tubulin in tension wood fibres of Populus
72 The significance of microfibril angle The relationship between MFA and axial stiffness according to Cave (1968) There is a 5 fold increase in stiffness when MFA shifts from 40 to 10 degrees
73 Corewood in a 125-year Old Tree Juvenile wood (large microfibril angle) Mature wood (small microfibril angle)
74 Corewood in a 25-year Old Tree Juvenile wood Mature wood
75 The Problem of High Microfibril Angle Corewood is too flexible to be used as high grade timber Any improvement that would reduce the amount of low grade timber would result in significant financial gain for the producer and result in more efficient use of forest land
76 Consequences for the Tree High microfibril angle in corewood makes young trees flexible and able to withstand high winds Only small reductions in angle may be feasible without affecting survivability These may, however, still give significant increases in the quality of corewood
77 Pitting and plasmodesmata
78 Formation of pits Aesculus hippocastanum vessel wall
79 Sorbus aucuparia Cambium pit fields
80 Pit fields in enlarging fibres
81 Pinus radiata Cambium pit fields
82 Pit fields in enlarging tracheids
83 Interference contract micrograph of a TLS through the radial wall of a tracheid of Pinus radiata
84 Pinus radiata Pit fields in radial walls of enlarging tracheids
85 Developing torus
86 Pinus radiata Beginning of formation of the torus and pit border
87 P. radiata Functional and aspirated bordered pits
88
89 Vessel pitting seen from the middle lamella in Aesculus
90 Vessel-vessel wall in Aesculus hippocastanum
91
92
93 Plasmodesmata in radial wall of cambium in Aesculus
94 Fibre-fibre pit in Aesculus
95 Pits in a tangential wall between ray parenchyma
96 Plasmodesmata in pit fields Absent: Vessels and tracheids (conifer and angiosperm) to any other cell type Present: Fibres to fibres and parenchyma Parenchyma to parenchyma and fibres
97 Plasmodesmata in developing vessel walls of hybrid aspen
98 Microtubules and pit formation
99 Tubulin in a developing Aesculus vessel
100 Tubulin in young vessel elements in Aesculus
101 Plasmodesmata in pit membranes in some members of the Rosaceae
102
103 Sorbus aucuparia RLS 2µm thick Plasmodesmata appear in section as black spots
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105 Pit fields in developing fibres of Sorbus
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113 Conclusions Microscopy reveals that cambial behaviour varies between species A knowledge of ultrastructural changes during differentiation of xylem is essential to understanding wood formation processes The cytoskeleton and plasmodesmata are important factors in the control of xylem differentiation Cambial behaviour ultimately governs wood structure and quality
114 The End
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