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Growth in Animals Determinate: Finite Size, Finite Shape, Number and Positions of Organs

Growth in Animals Determinate: Finite Size, Finite Shape, Number and Positions of Organs May have Larval Stages, Metamorphosis, but each is determinate Non-local regions of Cell Replication Growth in Plants Indeterminate: Final size, number of organs influenced by environment

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Growth in Animals Determinate: Finite Size, Finite Shape, Number and Positions of Organs

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  1. Growth in Animals Determinate: Finite Size, Finite Shape, Number and Positions of Organs May have Larval Stages, Metamorphosis, but each is determinate Non-local regions of Cell Replication Growth in Plants Indeterminate: Final size, number of organs influenced by environment Type of Organs influenced by environment Local regions of Cell Replication Repetitive Growth in Shoot Apex and in Radial/Lateral Meristems Continuous Growth in Root Apex

  2. Growth in Plants Local Increase in Cell Number (Cell Replication) Local Increase in Cell Size (Elongation)

  3. Introduction to Plant Growth Repetitive Production of Phytomeres: a Structural Module created by a Shoot Apical Meristem (SAM) Growth of shoots or roots in the Long or Axial Dimension is due to Shoot Apical Meristems (SAMs) and Root Apical Meristems (RAMs) These are regions at the apex that contain Stem Cells that replicate to form Daughter Cells while maintaining the Stem Cells As they are born and elongate, daughter cells of SAMs displace the SAMs upward and RAMs downward

  4. Region where axillary SAM will form Shoot SAM Root RAM Repetitive Growth From Shoot Apical Meristems (SAMs) Nonrepetitive Growth From Root Apical Meristems (RAMs) Growth from Apical Meristems and LDO Primordia

  5. Internode Phytomere Node Axillary SAM Apical End/ Apex /Medial Position Radial/Lateral / Lateral Position /Adaxial/ Top Long Axis = Axial Axillary Position/ Axil Lateral Determinate Organ (LDO) /Abaxial/ Bottom Basal End/ Base Axes in Plants

  6. SAM Daughter Cells form the Four Parts of the Phytomere Node: On a stem, the point of insertion of a Lateral Determinate Organ Lateral Determinate Organ (LDO): Leaf, Petal, etc. inserted into the Node. Some consider the LDO to be part of the Node. Axillary Shoot Apical Meristem (Axillary SAM): in the axil defined by the distal-most position between the Internode and LDO Internode: Portion of stem between Nodes The phytomere derives from the same group of founder cells In eudicots, the phytomere consists of a node, its axillary SAM, and the internode above the node In maize (a monocot), it includes the internode below the node

  7. Shoot Axillary Vegetative Shoot Apical Meristem (V-SAM) Florescence Phytomere Lateral Determinate Organ (LDO) Vegetative Phytomere

  8. 5 4 Phytomers 3 2 1 Vegetative Phytomers in Pea

  9. o 1 o 3 Active SAM o 2 o o 4 2 o 3 o 4 Inactive Axillary SAM Branching Shoots produced by Axillary Shoot Apical Meristems will usually be identical to the shoot on which they are borne, that is SAMs usually produce axillary SAMs of the same Phase

  10. A Case Study in Branching of Vegetative Shoots Monocot: Poaceae, Grass Family Genus: Phyllostachys, Bamboo Phyllostachys aureaCarrière ex A. & C. Rivière, Golden Bamboo, Fishpole Bamboo Introduced; Invasive; Spreads by underground stems = rhizomes; Does not Flower A ‘woody’ monocot, although not true wood (not made by vascular cambium)

  11. In this plant, SAMs know what order branch they are on Outgrowth of Both Axillary SAMs into Secondary Shoots Flat surface Flat surface Outgrowth of One of Two Axillary SAMs into a Tertiary Shoot Internode Node Two Axillary SAMs Both dormant Phytomere Outgrowth of One of Two Axillary SAMs into a Secondary Shoot Phyllostachys aureaCarrière ex A. & C. Rivière, Golden Bamboo, Fishpole Bamboo Phyllotaxy of Leaves of Primary Shoot = Distichous (2-Ranked; Alternate) [Do not have a term for ‘Two Adjacent Axillary SAMs per Node’]

  12. I NI NI NI N Internode Node Two Axillary SAMs Both dormant Phytomere Outgrowth of One of Two Axillary SAMs into a Secondary Shoot Phyllostachys aureaCarrière ex A. & C. Rivière, Golden Bamboo, Fishpole Bamboo Phyllotaxy of Leaves of Primary Shoot = Distichous (2-Ranked; Alternate) [Higher Magnification]

  13. A third axillary SAM at this node in the Primary Shoot! Remains dormant Phyllostachys aureaCarrière ex A. & C. Rivière, Golden Bamboo, Fishpole Bamboo Phyllotaxy of Leaves of Primary Shoot = Distichous (2-Ranked; Alternate) [Do not have a term for ‘Two (or Three!) Adjacent Axillary SAMs per Node’]

  14. Secondary Shoot Tertiary Shoot Two axillary SAMs per node in Secondary Shoots: One remains dormant. It is not random; they are on the same side! Tertiary Shoot One axillary SAM per node in Tertiary Shoots: remains dormant. If two axillary SAMs per node, then one grows out. Phyllostachys aureaCarrière ex A. & C. Rivière, Golden Bamboo, Fishpole Bamboo Phyllotaxy of Secondary and Tertiary Shoots = Distichous (2-Ranked; Alternate)

  15. Phase Transitions Induction of Flowering Converts Apical and Axillary AdultVeg-SAMs into one of several Reproductive SAMs JuvV-SAM AdultV-SAM Subflor-SAM Floresc-SAMFlower-SAM Large variation in position and complexity of the shoots that bear flowers: from axillary, solitary flowers to axillary and terminal, highly branched and complex shoots bearing many flowers

  16. Phase Transitions The SAM undergoes Phase Transitions: transitions in the Identity of the SAM as evidenced by the Identity of the Phytomeres (including the axillary SAMs) that the SAM produces Regardless of the Phase of the SAM, it still produces phytomeres composed of a node, internode, axillary SAM and (usually) a LDO, although their names may be phase specific Juvenile Vegetative Shoots - can branch from axillary JuvV-SAMs Adult Vegetative Shoots - can branch from axillary AdultV-SAMs Subflorescence Shoots - can branch from axillary Subflor-SAMs Florescence Shoots - cannot branch; axillary SAMs are Flower SAMs Flower Shoots - cannot branch; no axillary SAMs

  17. Vegetative Shoot Inflorescence Juvenile-Vegetative Adult-VegetativeSubflorescence FlorescenceFlower Shoot Shoot (Branched Shoot) (Floral Shoot)Shoot Lateral Juv Leaf Adult Leaf Cauline LeafBract S PSt C Determinate Organ (LDO) Axillary Shoot JuvV-SAM AdultV-SAM Subflor-SAMFlower-SAM Apical Meristem (axillary SAM) 1o Shoot ApicalJuvV-SAMAdultV-SAM Subflor-SAMFloresc-SAMFlower-SAM Meristem (SAM) 1o Full Series of Phase Tansitions with development of all LDOs and some axillary SAMs Terminal Inflorescences Only 3o 3o 3o 2o 2o 2o

  18. Vegetative Shoot Inflorescence Juvenile-Vegetative Adult-Vegetative FlorescenceFlower Shoot Shoot (Floral Shoot)Shoot Lateral Juv Leaf Adult Leaf Bract S PSt C Determinate Organ (LDO) Axillary Shoot JuvV-SAM AdultV-SAM Flower-SAM Apical Meristem (axillary SAM) 1o Shoot ApicalJuvV-SAMAdultV-SAM Floresc-SAMFlower-SAM Meristem (SAM) S P St C 1o Terminal Florescences; no Subflorescence Phase Plus Terminal Flower 3o 3o 2o 2o

  19. Vegetative Shoot Inflorescence Juvenile-Vegetative Adult-Vegetative FlorescenceFlower Shoot Shoot (Floral Shoot) Shoot Lateral Juv Leaf Adult Leaf Bract S PSt C Determinate Organ (LDO) Axillary Shoot JuvV-SAMFloresc-SAMFlower-SAM Apical Meristem (axillary SAM) 1o Shoot ApicalJuvV-SAMAdultV-SAM Floresc-SAMFlower-SAM Meristem (SAM) S P St C 1o Terminal and Axillary Florescences; no Subflorescence Phase Plus Terminal Flower 3o 3o 2o 2o

  20. Vegetative Shoot Inflorescence Juvenile-Vegetative Adult-VegetativeFlower Shoot Shoot Shoot S P St C Lateral Juv Leaf Adult Leaf S PSt C Determinate Organ (LDO) Axillary Shoot JuvV-SAMFlower-SAM Apical Meristem (axillary SAM) 1o Shoot ApicalJuvV-SAMAdultV-SAM Meristem (SAM) 1o Solitary, Axillary Flowers Plus Gradual Change in AdultV LDO 3o 3o 2o 2o

  21. Flower containing 4 phytomers Primary Florescence/ Floral Shoot (No Branching) Axillary SAMs are Flower-SAMs Florescence Phytomer Tertiary Inflorescence Subflorescence (Branched, Branchable) Subflorescence Phytomere Lateral Rosette Inflorescence Primary Inflorescence/ Reproductive Shoot Axillary SAMs are Subflor-SAMs Arabidopsis Vegetative Shoot Fig. 9.8A L&D Vegetative and Reproductive Structures and Phytomeres The Florescence is the apical, unbranched, part of an Inflorescence Warning: There are other nomenclatures for reproductive structures In Taxonomy, ‘Inflorescence’ usually includes Inflorescence, Florescence, and Flowers; for example, ‘Inflorescences axillary, solitary flowers’

  22. secondary paraclade 3 primary Types of Phytomeres: 1 = Basal [Vegetative] Rosette (same use as GG) 2 = Basal part of Flowering [Inflorescence] Stalk [the Subflorescence as used by GG] 3 = Florescence (same use as GG) Types of Branches containing …. Type 3 on Primary Shoot = Primary Florescence (Florescence on 1o shoot as used by GG) Type 3 Collectively = Coflorescence (no GG term) Type 2 and Type 1 = Paraclade (Higher-order Inflorescences as used by GG) Types 1+2+3 Collectively = Inflorescence 3 primary 3 2 2 rosette 1 An Alternative Nomenclature for Arabidopsis

  23. Vegetative Shoot Inflorescence Juvenile-Vegetative Adult-VegetativeSubflorescence FlorescenceFlower Shoot Shoot (Branched Shoot)(Floral Shoot) Shoot Lateral Juv Leaf Adult Leaf Cauline LeafNoneS PSt C Determinate Organ (LDO) Axillary Shoot Subflor-SAMNone Subflor-SAMFlower-SAM Apical Meristem AccessoryAccessory (SAM) Subflor-SAMSubflor-SAM 1o Shoot ApicalJuvV-SAMAdultV-SAM Subflor-SAMFloresc-SAM Meristem (SAM) 1o 2o 3o 2o 3o Arabidopsis violates several rules 1o 2o

  24. Vegetative Shoot Inflorescence Juvenile-Vegetative Adult-VegetativeFlower Shoot Shoot Shoot S P St C Lateral Juv Leaf Adult Leaf BractS P St C Determinate Organ (LDO) Axillary Shoot JuvV-SAMFloresc-SAM Apical Meristem (axillary SAM) 1o Shoot ApicalJuvV-SAMAdultV-SAM Meristem (SAM) 1o Axillary, not terminal, Florescences; no Subflorescence Phase Plus Gradual Change in AdultV LDO 3o 3o 2o 2o

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