The root and shoot both form in the dividing tissues at each end and grow like mirror reflections of each other.
The part of stem where buds grow is called node. The part between two buds is called internode. Since a bonsai is a miniaturization of a natural plant, you should shrink the internode. However, internode can grow overlong where the growth is vigorous. Such branches are known as coarse and vigorous branches.
We present here research findings on plant physiology and include their explanations for you to better understand the various operations in bonsai making. We summarize the general principles, and we will keep the explanations simple. However, the knowledge we include here is not required directly on a pragmatic basis when making a bonsai. Of course, there are many exceptions, for the diverse nature contains countless varieties of plants. Whatever the exceoptions there may be, you will learn to single them out after knowing the general principles. Perhaps you have already or will read professional instructions and heard or will hear excerpts' explanations before you start making a bonsai. But have you understood or will you understand the meaning of those explanations? If not, you may not be able to fully apply the knowledge, and you can easily forget what is written there. Also, when making a bonsai you should know whether your daily work fits the plant's physiological needs and how your work suits the plant's vital activity. Knowing these will add more fun to your bonsai making experience. In addition, when new problems arise and you solve them by trials and errors, the knowledge of the physiology of your plant can help you solve these puzzles. We hope the materials described here will become Ariadne's Thread in guiding your bonsai experience.
- Physiology Describes the Secret -
Principle of Plant's Shape
Unit of plant's shape --The Shoot A “shoot” botanically refers to the pair of stem and leaves growing on the stem. This set of two things forms the basic unit of a plant's shape. A plant begins with one shoot and grows more shoots as it develops. In this way a plant gradually forms its complex shape.
A plant consists of the shoots.
Shoot formed in the bud A shoot is first formed by the cell division and regeneration of the shoot apical meristem in the buds. It then gradually grows. In this process, cells in the shoot, i.e. the stem, and cells in the leaves and the buds (lateral buds or axillary buds) are formed. Newly formed stem, leaf, and lateral bud are compressed and folded in the bud.
A cabbage is a giant shoot grown directly from the bud
The development of the stem, leaf, and lateral bud during the shoot's growth As water is absorbed from outside the cells to the vacuoles inside the cells, it increases the volume of each cell and enlarges the cell. By the time cells mature, the volume of each cell will increase longitudinally by tens of times. Insufficient water in vacuoles and cells lowers the water pressure in each cell, which is unable to hold up the plant in the same way flat tires cannot hold up a vehicle. Therefore, not watering the plant on time will cause it to wither. As this process goes on, the folded stem, leaf, and lateral bud grow larger and stretch in the vertical direction.
Starting in the bud, shoot tissues including the stem, leaf, and bud develop gradually.
Two units of plant ― The correspondence between the shoot and root Besides the shoot, the root forms another unit in a plant's structure. The shoot mainly grows upwards. In contrast, the root mainly grows downwards. However, there are many exceptions. Some plants grow their roots above the ground, while other plants grow their shoots underground. The root and shoot both form in the dividing tissues at each end and grow like mirror reflections of each other. The cell division is the most active around both ends (cell division site). The cell division frequency decreases as one moves away from each end to the cell development site, where cells grow larger and stretch in vertical directions instead of dividing. Far away from each end, cells will not grow too large but they form distinct shapes according their functions. This is the cell maturation site.
The balance between the shoot and root
1.Balance in water The sizes of the root and shoot are determined by the balance between the amount of water absorbed in the root and that used for transpiration and photosynthesis in the shoot (mainly in the leaves). Water is mainly used for photosynthesis and cell growths. 2.Balance in nutrients Plants grow by cell divisions in the buds. For cell divisions to occur, nutrients such as minerals absorbed from the roots and carbohydrates produced by photosynthesis in the shoots (mainly in the leaves) are necessary. The root and shoot will grow larger until they can supply the plant with enough nutrients. 3.Balance in mechanics The root will grow thicker and larger from the taproot in the center until it can support the shoot and prevent it from leaning. Conversely, the shoot only grows as large as the root does.
For a plant growing in moist soil, its roots absorb plenty of water, so its shoots will keep growing. This will cause the plant to form coarse and vigorous branches. On the other hand, more branches and leaves will result in more transpiration from the shoot. So the roots will grow to balance. If you over-pruned or injured the roots during repotting, this sudden decrease in the amount of roots will disturb the balance between the roots and shoots. Then, the tree may defoliate or let part of the shoots wither to recover the balance. But failure to recover the previous balance will cause the entire plant to wither.
The growths of the shoot and root are balanced.
The root grows larger according to the needs of the shoot, and the shoot grows until the root can fully support it. The two parts are mutually dependent. The larger one part grows, so will the other part develop larger. Specifically, the balance between the shoot and root signifies the balance in the supply and demand of the following three factors in the plant. These factors determine the sizes of the shoot and root.
The leaves of deciduous trees fall in late autumn, and the leaves do not yet develop in early spring. So late autumn and early spring are the best seasons to root-prune plants in temperate zones. In these periods, leaves of evergreen trees are inactive, so the requirement of water and nutrients in the shoot lowers. Thus, the damage to the balance between the root and the shoot is minimal even if you prune the roots. Should you prune the roots, since the soil temperature has not dropped in late autumn or early spring, the roots are still active and are capable of healing the wounds after being pruned. For the same reasons as above, the best time to prune tropical plants is during their dormancy. If you prune your plant's roots at other times, you need to prune some leaves as well. You should also supply the plant with enough water after pruning. It should be mentioned that after pruning the roots on the left side of the plant, the leaves on that side may sometimes begin to wither to an extent. This suggests that specific part of the roots supply water and nutrients to specific part of the shoots.
Special Column
What is coarse and vigorous branch?
In order to reproduce a tree's natural shape, the bonsai tree should have a thick trunk that tapers upwards. The branches and leaves should be thinner and finer compared to the trunk, and they should grow amply. All these can be achieved by controlling and adjusting the root-shoot balance. When training your bonsai tree from a seeding or a young tree, you should first let the roots grow as large as possible in order to let the trunk grow thick. Since the amount of water and nutrients needed in the shoot is equal to that provided by the root, you should let the roots grow fully so that the shoot can fulfill its needs. In this stage, to ensure the roots will grow freely, you should either grow the plant in a large flowerpot or grow it in the ground. To make the shape of your bonsai, you need to repeatedly prune the stems, leaves, and buds on the shoot. The bonsai's shape is made by pruning the shoots and adjusting the balance between the root and the shoot. By pruning large amount of shoot under an existing root-shoot balance, you reduce more rapidly the consumption of water and nutrients in the shoot than the supply by the root. Under such stimulation, the extra water and nutrients supplied by the root will flow to the hitherto dormant lateral buds and activate their growths. Specifically, when stems are pruned, the buds will of course be cut off as well. This will stop the growth of the stems in the vertical direction. Water and nutrients supplied by the root to the buds will then flow to the lateral buds and make them grow. Therefore, after you prune the stems above the lateral buds, new shoots will branch out near the cuts.
If the lateral buds have already developed into a new shoot, the new shoot will diverge more. The stems on the new shoot will be thinner than those on the old shoots. You can make use of this and prune the trunk and branches to make tapers. You can also make bends in the trunk and branches. In addition, you can apply this when pruning the coarse and vigorous branches to grow new branches.
After each leaf pruning, water and nutrients that has been previously supplied to the leaves will now flow to the lateral buds. Just like in stem pruning, the lateral buds on the stems will be activated and develop into a new shoot and keep growing. Pruning all the leaves once will cause huge imbalance between the water-and-nutrient supply by the root and the demand for it in the shoot. As a result, new shoots will develop in most abundance. For example, when you prune all the leaves after the buds have developed into a shoot and the leaves have matured, the tree will grow equal amount of leaves to restore the balance. This then activates the hitherto dormant lateral buds that will grow into new shoots. In that case, the tree in fact has completed a two-year-growth within one year. So you have reduced the time and have completed within one year a two-year training of your bonsai tree. If you can prune three times in one year, then you will accomplish three-year's training. In bud pruning, the shoots will stop growing like in stem pruning, and the lateral buds will be activated and grow into new shoot. Depending on the tree species, or if the tree is old or the momentum of its growth is low, some lateral buds may not be easily activated after stem pruning. Also, after leaf-pruning conifer trees, the leaf shape can deteriorate or the trees can easily wither. As you will probably spend the most energy in bud-pruning, the bud-pruning is used mainly in the above-mentioned trees, or uncommon trees, or deciduous trees whose branches and leaves you want to beautify.
Buds are essential for training new branches and making a bonsai. Buds grow out at the end or in the middle of a shoot. A bud growing at the end of a shoot is called apical bud, terminal bud, or end bud. That which grows in the middle of a shoot is called lateral bud, or axillary bud for it grows at the same place as the leaves do. The lateral buds grow to produce new shoots and become apical buds, so new shoots are also called lateral branches. In addition, buds can sometimes grow out on trunk, roots, and leaves. These buds are called adventitious buds. Flowering plants have flower buds or floral buds of slightly different shapes. So to differentiate from them, buds are sometimes called leaf buds or foliar buds.
In naturally growing trees, the shoots including the stems, branches and leaves develop larger upwards and stretch outwards as they grow. The opposite should happen in a bonsai, the tree must remain small while it grows. So you must prune the stems, branches, and leaves that have grown upwards or outwards to make a bonsai. Then relatively more water and nutrients are supplied by the roots, and new shoots will grow. Water and nutrients now are supplied to the lateral buds on the inner side of the tree. These buds will be activated and grow in horizontal direction. As the lateral buds on the inner side grow, you can add more branches and leaves to the bonsai tree while keeping its original size. In addition, as the growth period is short, new branches and leaves are thin and small. You should observe the most upward and outward stretches at the shoot tips, i.e. the buds on the outside of the tree and on the coarse and vigorous branches. So you need to prune these parts often. In doing so, water and nutrients supplied by the roots will be transferred to other parts of the tree, which in turn accelerates the growth of the branches and leaves that have been growing slowly. Thus, the tree as whole has a balanced growth. Therefore, reducing the bonsai tree's size is intimately related to reproducing the tree's natural beauty.
The above-mentioned tips will only work when sufficient water and nutrients are supplied by the roots. According to botanical research findings, the size and quantity of leaves and the height of a tree in nature are determined by water and nutrients absorbed in soil and the total energy obtained from photosynthesis. You need to prune the roots to ensure they supply enough water and nutrients. The roots absorb water and nutrients in soil by root hair at the root tips. Root hair has a rather short life span, and new root hair constantly replaces old withering one. The most important factor in increasing water and nutrient absorption is not to have thick and long roots but to increase their total amount. Indeed, more roots can absorb more water and nutrients. For naturally growing trees, large and thick roots such as the taproots are necessary for supporting the shoots. But a bonsai tree does not need thick roots including the taproot. On the contrary, the taproot will inhibit the growth of finer roots. So during root pruning, you should focus on pruning the taproot and the thick roots. Like the shoots, new lateral roots will grow out after root pruning, which increases the total number of roots. More roots will provide more water and nutrients to the tree, which is advantageous to the development of lateral buds after shoot pruning. On the other hand, without root pruning there will be few roots, and so the shoots will be poor. Therefore, root pruning is indispensable to achieving the effect of shoot pruning.
Special Column
Plant hormone
Pruning makes the lateral buds grow, so the shoots and roots can grow to adjust their mutual balance. All these activities are due to the effects of plant hormones such as auxins and cytokinins. Plant hormores are chemicals that regulate the growth of the plant. There are several categories of them, but until now many aspects of these hormones have not been understood. Auxins and cytokins are not just two specific hormones, but they refer to two families of chemicals. They have various effects, and their main effect is to promote cell divisions. The most typical one is in forming the scab-looking knobby calluses which cover the wounds after the tree is cut or wounded in order to keep out germs and prevent tree juices from flowing out. Mutual effects of auxins and cytokinins in the growth of the shoots and roots Auxins are primarily synthesized in the apical buds in the shoots. They reach the roots through phloem in the stem and trunk. When the auxins concentration in the roots has reached a certain threshold, many lateral roots will grow out. Thus, a plant with active apical buds will form many branching roots. On the other hand, auxins can also inhibit the roots from growing larger. According to the results, as the auxins concentration increases, the plant grow out many tiny roots. Cytokinins are mainly synthesized in the apical meristem in the root tips. They are transported from the roots to the shoots with water and nutrients through xylem in the stem and trunk. Cytokinins will accumulate in the apical buds. This boosts the vigor of cell divisions there and thereby increases the vitality of the shoots and leaves. Therefore, it is correct to say more roots or more apical meristems poduce more cytokinins, and so shoots will grow more vigorously. Moreover, vigorously growing shoots will increase the amount of auxins synthesized in the apical buds and sent to the roots, which further boosts cytokinins synthesis there. These findings are scientific explanations of the fact that the trees with ample roots have abundant shoots. They also explain why plants stop growing in winter. As the soil temperature drops in winter, the roots become less active and so cytokinins synthesis decreases. Consequently, cell divisions stop in the apical buds.
Special Column
The reason why the lateral buds grow after being pruned――Apical dominance
Usually the apical buds alone are stimulated into growing, whereas the growth of all lateral buds below them is suppressed. This is called apical dominance. Shoots do not grow in lateral direction but grow outwards. Pruning is to cut off apical buds so as to remove the apical dominance. We consider the following hypotheses to explain the effects of auxins, cytokinins, and apical dominance. Dormant lateral buds will grow immediately after being applied cytokinins. Also, if we spray cytokinins to the entire tree, germination of the lateral buds will be evidently be boosted. Thus we can conclude that the apical dominance occurs because the cytokinins concentration in the apical buds is too high. In other words, the cytokinins concentration is the highest in the roots where they are synthesized, their concentration gradually lowers as they are transported upwards through xylem. In the topmost apical buds where cytokinins accumulate, their concentration is especially high. Auxins produced in the apical buds promote cells' absorption of cytokinins, so cytokinins accumulate especially in the apical buds with active auxins production. In contrast, cytokinins do not accumulate in the lateral buds, so the growth of the lateral buds is suppressed. The optimal auxins concentrations for promoting the growth of the apical buds and lateral buds differ. We have seen high concentration of auxins promotes the growth of the apical buds but suppress that of the lateral buds. We may consider a second hypothesis that lower auxins concentration is needed to promote the growth of the lateral buds. Indeed, if we prune the apical buds and place agar gel containing auxins on the wounds immediately after the lateral buds grow out, we will observe the growth of the lateral buds are suppressed. This confirms our hypothesis. In addition, the growth of the lateral buds after pruning the apical buds is the results of both the decrease in auxins and the increase in cytokinins. Besides the above mentioned plant hormones, other new chemicals have been discovered and identified more recently.
All plants develop from a single shoot and grow new shoots, on which new shoots grow out. As the shoot ends develop and branch for several times, the plant grows bigger. Meanwhile, the stems grow thicker to support the increased weight of the plant, and to transport more water and nutrients. Similarly, the roots keep growing. For the shoots and roots to grow thicker, cell reproduction must occur in the horizontal direction. This is called the secondary growth. It occurs in the cell division tissue known as the vascular cambium in the stems. It should be noted that some plants do not have secondary growths. But almost all plants used for bonsai have secondary growth. As the cells develop in vascular cambium, their cell walls thicken. After the cell die, their cell walls remain and harden, and become firm supportive tissue. Grass has short secondary growth, whereas trees have extended secondary growths. In this way trees make most of their stems, which are known as wood. Secondary growth occurs both in shoots and roots.
Water, nitrogen, and inorganic minerals in soil are absorbed by root hair and then transferred upwards through the vessels in xylem to the main root, trunk, branches, and eventually leaves and buds at the shoot ends. Water is broken down into oxygen and hydrogen by the energy in sunlight in the leaves, and photosynthesis produces carbohydrates such as sugar and starch by combining the hydrogen with the carbon dioxide absorbed from the stoma in the leaves. About 20% of water is not used in this process, instead it evaporates from the stoma in the leaves.
A plant is composed of cells and fibers. Cells support various vital life activities of the plant by cell divisions. Fibers build and shape the cell walls surrounding cells. Indispensable to all these life activities are carbohydrates. There are two types of cell activities. One concerns the growth of the plant, i.e the enlargement of its size, and the other its reproduction. For the plant to grow larger, the ends of the stems and branches will bud and the tree will grow taller. The branches will grow longer and the leaves will grow denser. As the roots penetrate deeper into the soil, the trunk and branches will grow thicker. The tree's wounds are also healing as it repairs itself. To produce offspring, the tree will flower and eventually bear fruit. Fibers make up the cell walls that surround the cells and make them rigid. In their development, cells first divide to produce more cells. Then the cell walls thicken to fix the plant tissues. The support provided by the cell walls enable the vessels transporting water and nutrients to maintain their shapes. The leaves and branches can keep their shapes for the same reason. Indeed, a plant can support itself on the ground because of the cell walls. Most tissues in the trunk and branches are composed of thick layers of cell walls left by dead cells. Carbohydrates are the building materials for cells and fibers. Carbohydrates combine with each other to form fibers. They react with nitrogen absorbed from roots into amino acids and proteins, which construct the shapes of plant cells. All above-mentioned processes require carbohydrates that are produced in the leaves and stored in the trunk and roots.
For trees growing in nature, the stem size is determined by the amount of water and nutrients transported through the vessels in xylem and that of carbohydrates transported through sieve tubes in phloem. In addition, the stem size should have enough wood to support the tree's weight. For a bonsai plant, you may neglect the weight issue, so the other two factors are more important. Here are some findings from the botanical research.
When a single stem branches into two, the cross section area of the stem before branching is more or less the same as the sum of the cross section areas of the two branches.
The cross section area of a stem is proportional to the total amount of leaves on the stem up to its end.
Simply put, the more a shoot branches out upwards, the thicker the shoot grows. So if you want to thicken a shoot, you should increase the number of branching at that shoot and increase the amount of leaves on it. For example, to thicken trunk while making “kokejun” (taper), you can just let more branches grow out of the trunk and let the leaves grow denser. Meanwhile, you should gradually reduce the amount of branches and leaves on the shoots as you move up the trunk, and cut off the topmost branches and leaves. As shoots grow via branching, this method can be applied to any part of a tree.
Carbohydrates produced in the leaves move to the branches, stems, and roots through sieve tubes in phloem. Eventually, they are transported to the buds and root tips where cell division is active, or to the reserve tissue in the roots and the stems to be stored. The opposite can also happen as carbohydrates move from the storage tissues in the stems and roots to the leaves and buds. Water, carbohydrates, and other nutrients are transported through xylem and phloem via changes in osmotic pressure and the capillary phenomenon.