Plant growth and development The need to explain tropisms: re-direction of growth in response to light, PHOTOTROPISM gravity, GEOTROPISM touch, THIGMOTROPISM The need to explain patterns of development: production of flowers, PHOTOMORPHOGENESIS development of fruits, senescence of foliage, response to wounds Phototropism The cress shoot was grown in unidirectional light from the right. There is a rapid curving response in the apical region which moves down the stem. Straightening of the upper stem in the later stages of the sequence makes the shoot appear to stop curving, but close examination of the lower stem shows there is still a response towards the light source. 180 frames filmed over 4 hours at 0.013 f.p.s. http://www-users.york.ac.uk/~drf1/tropism/tropisms.htm Negative gravitropism in a shoot A young, vertically growing, sunflower seedling shoot placed in a horizontal position. The shoot curves until it is once again growing vertically. This gravitropic response is first observed in the apical region of the shoot. Rapid curving at the tip progresses along the whole stem. The rate of curvature is not necessarily constant along the stem and is complicated by subsequent straightening of curved areas known as autotropism. 180 frames filmed over 5 hours at 0.01 f.p.s. Experimental analysis of shoot phototropism A sequence of experiments started by the Darwins and continued by Boysen-Jensen Fig. 33.1C Components of experiments An experiment has: A statement predicting alternative responses: 1. Hypothesis 2. Treatment “If this is done that will happen otherwise it will not.” A specific, designed, manipulation 3. Measurement 4. Control sufficiently accurate to detect response to the treatment The same measurement is made but the treatment is not applied. This provides the essential contrast. 5. Replication Enables the degree of response to be defined and helps to protect against obtaining results by chance And must be 6. Repeated Required to establish the degree of certainty that can be attributed to a result, e.g., repetition with the same and different species The initial observation The first experiment to remove the effect 1. Hypothesis 2. Treatment 3. Measurement 4. Control 5. Replication And must be 6. Repeated Deficiencies? 1. Hypothesis The initial observation Expt 1 Expt 2 Expt 3 2. Treatment 3. Measurement Expts 2 and 3 4. Control 5. Replication And must be 6. Repeated Does the postulate change for Expt 2 and 3? What are the improvements over the first experiment? What are the treatments and what the controls in Expt2 and Expt3? What type of an experiment is this? The initial observation 1. Hypothesis 2. Treatment Expt4 3. Measurement 4. Control 5. Replication And must be 6. Repeated Boysen-Jensen’s experiment The initial observation 1. Hypothesis 2. Treatment 3. Measurement 4. Control 5. Replication And must be 6. Repeated Has the postulate changed from that of Expt 4? What are the improvements over Expt4 ? Which is the treatments and what are the controls? Went’s experiments Is this really a control? Fig. 33.1D Chemicals are produced in small quantities that change the rate at which growth takes place and/or the types of cells that are produced. These chemicals are usually produced by meristematic tissue and are actively transported from that tissue. The chemicals influence the development of cells according to the concentration that accumulates in the developing cells. Plant growth substances, the text book calls them hormones Five plant growth substances and their functions Major Functions Where Produced Stimulates cell elongation; involved in phototropism, gravitropism, apical domincance, and vascular differentiation; stimulates ethylene synthesis and induces adventitious roots on cuttings Meristems of apical buds, embryo of seed, young leaves Cytokinin Stimulates cell division, reverse apical dominance, involved in shoot growth, delay leaf sequence Synthesized in roots and transported to other organs Ethylene Stimulates fruit ripening, leaf and flower senescence, and abscission Tissues of ripening fruits, nodes of stems, senescent leaves and flowers Abscisic Acid Inhibits growth, stimulates stomatal closure, maintains dormancy Leaves, stems, green fruit Stimulates shoot elongation, stimulates bolting and flowering in biennials, regulates production of hydrolytic enzymes in grains Meristems of apical buds and roots, young leaves, embryo Auxin Gibberellin Auxin and phototropism Auxin, indole acetic acid (IAA), is transported downwards from the shoot apex. In a seedling exposed to light from the side the concentration of IAA is greater on the shaded side of the shoot Fig. 33.1B Results of experiments that applied IAA Fig. 33.3B 1. The chemical may have different effects at different concentrations 2. It can affect different tissues differently Polar transport of auxin Transport at ~1 cm/hr implies active transport Picks up a hydrogen ion at the acid wall environment Passes across membrane as a neutral molecule Gives off the H+ into the cell which induces the proton pump Auxin can only exit the cell at its basal end where there are specific carrier proteins The acid growth hypothesis Gibberellins •Reverses dwarfism – the first discovery of gibberellin •Seed Germination--Barley de novo amylase synthesis (Varner 1964) •Can cause bolting in biennials •Control of sex expression •Can enhance fruit growth – e.g., seedless grapes •Delays Senescence Gibberellins act in the elongation of intact plants as opposed to stem section elongation by auxin. Much research on plant gibberellins has been possible due to gibberellin sensitive mutants. They have adequate levels of GA1 (the GA species most likely to be responsible for stem elongation) but can not respond to it. This may be due to lack of receptor protiens. Transport is non polar, has been found in both the transpiration and translocation stream. It can occur more rapidly, 5 cm/hr, than auxin There are many gibberellins – closely related chemically Bioassay Using an organism or other living material to determine the level of an environmental condition 1. Establish pattern of response under standardized conditions 2. Use the established pattern in analysis of treatments Advantages: cost, useful where the treatment has not been precisely defined yet, e.g., substances ‘like’ gibberellin. The effect of day length on flowering Autumn flowering plants, e.g., chrysanthemums Summer flowering plants, e.g., iris Fig. 33.11 Flowering response can be manipulated by short periods of red or far-red radiation applied during the dark period of a long night regime Fig. 33.12A Phytochrome The control of flowering is determined by a substance called phytochrome that exists in two forms. 660 nm 730 nm All plants contain phytochrome – but they may respond differently to the relative amounts of the two forms Phytochrome is involved in other plant growth process in addition to flowering Control through the relative amounts of different Plant Growth Substances Abscissic acid as a growth inhibitor gibberellin as a promoter For cells growing in culture: Cytokinins added have no effect on their own. Cytokinins plus auxin cause cells to divivde. If the concentrations are about equal the cells continue to grow and form a callus but there is no cell differentiation. If there is more cytokinin than auxin then shoot buds develop. If there is more auxin than cytokinin then roots develop Are plant growth substances ‘hormones’? A hormone is a regulatory chemical that travels in the blood from its production site and affects other sites in the body often at some distance. Hormones are made and secreted by organs called endocrine glands. This hormone concept as developed for animals has some distinct differences from what we know of the production, distribution and function of plant growth substances. 1. Transport in plants is very different from that in animals. Its is frequently polar 2. The range of plant growth substances that we know are produced and possibly distributed in different ways 3. Plants no equivalent to the central nervous system that integrates and co-ordinates physiological activities. Sections you need to have read 33.1 through 33.12 Courses that deal with this topic Botany 371/372 Plant physiology laboratory