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Section 9: Branching of Shoot Systems

      Conventional cauline branching originates from lateral/axillary buds. When a lateral bud expands, it becomes a lateral branch or shoot. Subsequent branching gives rise to ranks or orders of branches. These are referred to as primary branches, secondary branches, tertiary branches, and so on.

      Many species of plants possess more than a single axillary bud in their leaf axils. The additional buds are called accessory or supernumerary buds. Such buds can have specific patterns of initiation and can differ from species to species. The developmental fates of each bud within a leaf axil may be quite specialized.

serial_buds.png

      Accessory buds usually vary in size and in most cases the larger buds will continue to develop and the smaller ones will remain dormant.

      Lateral buds may also be present in the axils of cotyledons and are sometimes located in the hypocotyl region of the embryo. Axillary buds of cotyledons provide a means of survival if the epicotyl portion of certain plants are removed by herbivores, etc. Once the apical meristem is excised, the cotyledon axillary bud begins to expand and develop. Subcotyledonary buds can be found on the noxious weed Convolvulous arvensis or bindweed. These buds allow the plant to produce new shoots from roots and to survive, even when older shoots are removed by herbivores and avid gardeners.

Lateral Branching
Lateral Branching

      One common form of branching, not involving buds, is dichotomous branching. This occurs when a division of an apical meristem forms into two equal parts. The classic example of dichotomous branching is found in the macroscopic brown alga Dictyota, but dichotomous branching is a common phenomenon on Marchantia and Psilotum as well as a host of other plants. Some plants exhibit this form of branching in their root systems also. This type of branching is not limited to the "lower" classes of plants but is found across the board of the land group classifications.

Dichotomous Branching
Dichotomous Branching

      Unequal development of shoots derived from dichotomously divided meristems results in anisotomous branches (of unequal length) as seen in some Lycopodium species. Others exhibit isotomous dichotomous branching patterns, where each of the branches of the dichotomy grow to equal lengths.

 

Equisetum

      Lateral branching of Equisetum occurs at the nodes but cannot be considered strictly axillary due to the unique morphology of the leaves which are appressed to the stem. The leaves are united near their bases by a membraneous sheath that surrounds the entire stem and lateral branches must forge their way through this tissue during the course of their development. This occurs at alternating radii of the leaves, resulting in a whorl of lateral branches.

 

Equisetum
Equisetum

Monopodial vs. Sympodial Branching

      Monopodial branching is a pattern of branching in which there is one main shoot with lateral branches emerging from it. Many firs and spruces exhibit monopodial branching patterns, therefore the Christmas tree image is a good example of monopodial branching.

      Sympodial branching occurs when the apical meristem of the main shoot is determinant in its growth pattern. Some time later one or more lateral buds expand and become the new "leading shoots." This type of developmental pattern results in a highly branched structure.

Monopodial vs. Sympodial
Monopodial vs. Sympodial
monopodial_sympodial

 

Acrotonic vs. Basitonic Branching Patterns

      These terms describe the developmental growth of monopodial shoots. Acrotonic branching occurs when a series of lateral branches emerge from the main shoot. This shoot then undergoes expansion that lengthens the shoot. The lateral branches nearest the apex elongate earlier, and grow for a longer period of time than the lateral branches further from the apex. The resulting form has the longest branches up near the terminus of the shoot.

      Basitonic branching results when the lateral buds elongate in the order in which they were produced, thus the branches near to the base of the plant have undergone longer periods of expansion than those nearest the apex. This type of branching is seen in many shrubs and is responsible for their lower profiles.

      Branching patterns in many plants are a result of environmental factors. For instance, trees growing in the shade may have fewer, flabellate (flat) lateral branches that have more orders of branches as a means to collect as much sunlight as possible. Likewise, trees that are exposed to a great deal of sunlight are often conical in shape and have many lateral branches with only 3-5 orders.

acrotonic basitonic

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