Auxins

Nature of Auxins
The term auxin is derived from the Greek word auxein which means to grow. Compounds are generally considered auxins if they can be characterized by their ability to induce cell elongation in stems and otherwise resemble indoleacetic acid (the first auxin isolated) in physiological activity. Auxins usually affect other processes in addition to cell elongation of stem cells but this characteristic is considered critical of all auxins and thus "helps" define the hormone (Arteca, 1996; Mauseth, 1991; Raven, 1992; Salisbury and Ross, 1992).

History of Auxins and Pioneering Experiments
Auxins were the first plant hormones discovered. Charles Darwin was among the first scientists to dabble in plant hormone research. In his book "The Power of Movement in Plants" presented in 1880, he first describes the effects of light on movement of canary grass (Phalaris canariensis) coleoptiles. The coleoptile is a specialized leaf originating from the first node which sheaths the epicotyl in the plants seedling stage protecting it until it emerges from the ground. When unidirectional light shines on the coleoptile, it bends in the direction of the light. If the tip of the coleoptile was covered with aluminum foil, no bending would occur towards the unidirectional light. However if the tip of the coleoptile was left uncovered but the portion just below the tip was covered, exposure to unidirectional light resulted in curvature toward the light. Darwin's experiment suggested that the tip of the coleoptile was the tissue responsible for perceiving the light and producing some signal which was transported to the lower part of the coleoptile where the physiological response of bending occurred. He then cut off the tip of the coleoptile and exposed the rest of the coleoptile to unidirectional light to see if curving occurred. Curvature did not occur confirming the results of his first experiment (Darwin, 1880). It was in 1885 that Salkowski discovered indole-3-acetic acid (IAA) in fermentation media (Salkowski, 1885). The isolation of the same product from plant tissues would not be found in plant tissues for almost 50 years. IAA is the major auxin involved in many of the physiological processes in plants (Arteca, 1996). In 1907, Fitting studied the effect of making incisions on either the light or dark side of the plant. His results were aimed at understanding if translocation of the signal occurred on a particular side of the plant but his results were inconclusive because the signal was capable of crossing or going around the incision (Fitting, 1907). In 1913, Boysen-Jensen modified Fritting's experiment by inserting pieces of mica to block the transport of the signal and showed that transport of auxin toward the base occurs on the dark side of the plant as opposed to the side exposed to the unidirectional light (Boysen-Jensen, 1913). In 1918, Paal confirmed Boysen-Jensen's results by cutting off coleoptile tips in the dark, exposing only the tips to the light, replacing the coleoptile tips on the plant but off centered to one side or the other. Results showed that whichever side was exposed to the coleoptile, curvature occurred toward the other side (Paal, 1918). Soding was the next scientist to extend auxin research by extending on Paal's idea. He showed that if tips were cut off there was a reduction in growth but if they were cut off and then replaced growth continued to occur (Soding, 1925). In 1926, a graduate student from Holland by the name of Fritz Went published a report describing how he isolated a plant growth substance by placing agar blocks under coleoptile tips for a period of time then removing them and placing them on decapitated Avena stems (Went, 1926). After placement of the agar, the stems resumed growth (see below). In 1928, Went developed a method of quantifying this plant growth substance. His results suggested that the curvatures of stems were proportional to the amount of growth substance in the agar (Went, 1928). This test was called the avena curvature test.

Much of our current knowledge of auxin was obtained from its applications. Went's work had a great influence in stimulating plant growth substance research. He is often credited with dubbing the term auxin but it was actually Kogl and Haagen-Smit who purified the compound auxentriolic acid (auxin A) from human urine in 1931 (Kogl and Haagen-Smit, 1931). Later Kogl isolated other compounds from urine which were similar in structure and function to auxin A, one of which was indole-3 acetic acid (IAA) initially discovered by Salkowski in 1985. In 1954 a committee of plant physiologists was set up to characterize the group auxins. The term comes from the Greek auxein meaning "to grow." Compounds are generally considered auxins if they are synthesized by the plant and are substances which share similar activity to IAA (the first auxin to be isolated from plants) (Arteca, 1996; Davies, 1995).

Biosynthesis and Metabolism of Auxin
IAA is chemically similar to the amino acid tryptophan which is generally accepted to be the molecule from which IAA is derived. Three mechanisms have been suggested to explain this conversion: Tryptophan is converted to indolepyruvic acid through a transamination reaction. Indolepyruvic acid is then converted to indoleacetaldehyde by a decarboxylation reaction. The final step involves oxidation of indoleacetaldehyde resulting in indoleacetic acid. Tryptophan undergoes decarboxylation resulting in tryptamine. Tryptamine is then oxidized and deaminated to produce indoleacetaldehyde. This molecule is further oxidized to produce indoleacetic acid. As recently as 1991, this 3rd mechanism has evolved. IAA can be produced via a tryptophan-independent mechanism. This mechanism is poorly understood, but has been proven using trp(-) mutants. Other experiments have shown that, in some plants, this mechanism is actually the preferred mechanism of IAA biosynthesis.

The enzymes responsible for the biosynthesis of IAA are most active in young tissues such as shoot apical meristems and growing leaves and fruits. The same tissues are the locations where the highest concentrations of IAA are found. One way plants can control the amount of IAA present in tissues at a particular time is by controlling the biosynthesis of the hormone. Another control mechanism involves the production of conjugates which are, in simple terms, molecules which resemble the hormone but are inactive. The formation of conjugates may be a mechanism of storing and transporting the active hormone. Conjugates can be formed from IAA via hydrolase enzymes. Conjugates can be rapidly activated by environmental stimuli signaling a quick hormonal response. Degradation of auxin is the final method of controlling auxin levels. This process also has two proposed mechanisms outlined below: The oxidation of IAA by oxygen resulting in the loss of the carboxyl group and 3-methyleneoxindole as the major breakdown product. IAA oxidase is the enzyme which catalyzes this activity. Conjugates of IAA and synthetic auxins such as 2,4-D can not be destroyed by this activity. C-2 of the heterocyclic ring may be oxidized resulting in oxindole-3-acetic acid. C-3 may be oxidized in addition to C-2 resulting in dioxindole-3-acetic acid. The mechanisms by which biosynthesis and degradation of auxin molecules occur are important to future agricultural applications. Information regarding auxin metabolism will most likely lead to genetic and chemical manipulation of endogenous hormone levels resulting in desirable growth and differentiation of important crop species. Ultimately, the possibility exists to regulate plant growth without the use of hazardous herbicides and fertilizers (Davies, 1995; Salisbury and Ross, 1992).

Functions of Auxin
The following are some of the responses that auxin is known to cause (Davies, 1995; Mauseth, 1991; Raven, 1992; Salisbury and Ross, 1992).

  • Stimulates cell elongation
  • Stimulates cell division in the cambium and, in combination with cytokinins in tissue culture
  • Stimulates differentiation of phloem and xylem
  • Stimulates root initiation on stem cuttings and lateral root development in tissue culture
  • Stimulates root initiation on stem cuttings and lateral root development in tissue culture
  • Mediates the tropistic response of bending in response to gravity and light
  • The auxin supply from the apical bud suppresses growth of lateral buds
  • Delays leaf senescence
  • Can inhibit or promote (via ethylene stimulation) leaf and fruit abscission
  • Can induce fruit setting and growth in some plants
  • Involved in assimilate movement toward auxin possibly by an effect on phloem transport
  • Delays fruit ripening
  • Promotes flowering in Bromeliads
  • Stimulates growth of flower parts
  • Promotes (via ethylene production) femaleness in dioecious flowers
  • Stimulates the production of ethylene at high concentrations

References
Arteca, R. (1996). Plant Growth Substances: Principles and Applications. New York: Chapman & Hall.

 

Addicott, F. T., Lyon, J. L., Ohkuma, K., Thiessen, W. E., Carns, H. R., Smith, O. E., Cornforth, J. W., Milborrow, B. V., Ryback, G., and Wareing, P. F. (1968). "Abscisic acid: A new name for abscisin II (dormin)". Science 159:1493.

Bandurski, R. S., Cohen, J. D., Slovin, J., and Reinecke, D. M. (1995). "Auxin biosynthesis and metabolism". Plant Hormones: Physiology, Biochemistry and Molecular Biology. Dordrecht: Kluwer. pp. 39-65.

Boysen-Jensen, P. (1913). "Uber die Leitung des phototropischen Reizes in der Avenakoleoptile". Ber. Deut. Bot. Ges. 31:559-566.

Brian, P. W., Elson, G. W., Hemming, H.G., and Radley, M. (1954). "The plant-growth promoting properties of gibberellic acid, a metabolic product of the fungus Gibberella fujikuroi". J. Sci. Food. Agr. 5:602-612.

Crocker, W., Hitchcock, A. E., and Zimmerman, P. W., (1935). "Similarities in the effects of ethylene and the plant auxins". Contrib. Boyce Thompson Inst. 7:231-248.

Darwin, C. R. (1880). The Power of Movement in Plants. London: Murray.

Davies, P. J. (1995). Plant Hormones: Physiology, Biochemistry and Molecular Biology. Dordrecht: Kluwer.

Doubt, S. L. (1917). "The response of plants to illuminating gas". Bot. Gaz. 63:209-224.

du Monceau, D. (1758). La Physique des arbres. Volume I.

Fitting, H. (1910). "Weitere entwicklungsphysiologische Untersuchungen an Orchideenbluten". Zeitschr. Bot. 2:225-267.

Fitting, H. (1907). "Die Leitung Tropistischer Reize in parallelotropen Pflanzenteilen", Jahrb. Wiss. Bot. 44:177-253.

Gane, R. (1934). "Production of ethylene by some ripening fruits". Nature 134:1008.

Haagen-Smit, A. J., Dandliker, W. B., Wittwer, S. H., and Murneek, A. E. (1946). "Isolation of 3-indoleacetic acid from immature corn kernels". Amer. J. Bot. 33:118-120.

Haberlandt, G. (1913). "Zur Physiologie der Zellteilung". Sitzber. K. Preuss. Akad. Wiss. 318.

Hall, R. H. and deRopp, R. S. (1955). "Formatin of 6-furfurylaminopurine from DNA breakdwon products". J. Am. Chem. Soc. 77:6400.

Hori, S. (1898). "Some observations on 'bakanae' disease of the rice plant". Mem. Agric. Res. Sta. (Tokyo) 12:110-119.

Jablonski, J. R. and Skoog, F. (1954). "Cell enlargement and cell division in excised tobacco pith tissue". Physiol. Plant. 7:16.

Kogl, F. and Haagen-Smit, A. J. (1931). "Uber die Chemie des Wuchsstoffs K. Akad. Wetenschap. Amsterdam". Proc. Sect. Sci. 34:1411-1416.

Kurosawa, E. (1926). "Experimental studies on the nature of the substance secreted by the 'bakanae' fungus". Nat. Hist. Soc. Formosa 16:213-227.

Lang, A. (1970). "Gibberellins: Structure and Metabolism". Annu. Rev. Plant. Physiol. 21:537-570.

Letham, D. S. (1963). "Zeatin, a factor inducing cell division isolated from zea mays". Life Sci. 2:569-573.

MacMillan, J. and Takahashi, N. (1968). "Proposed procedure for the allocation of trivial names to the gibberellins". Nature 217:170-171.

Mauseth, J. D. (1991). Botany: An Introduction to Plant Biology. Philadelphia: Saunders. pp. 348-415.

McGaw, B. A. (1995). "Cytokinin biosynthesis and metabolism". Plant Hormones: Physiology, Biochemistry and Molecular Biology. Dordrecht: Kluwer. pp.98-117.

McKeon, T. A., Fernandez-Maculet, J. C. and Yang, S. F. (1995). "Biosynthesis and metabolism of ethylene". Plant Hormones: Physiology, Biochemistry and Molecular Biology. Dordrecht: Kluwer. pp. 118-139.

Miller, C. O. (1961). "A kinetin-like compound in maize". Proc. Natl. Acad. Sci. USA. 47:170-174.

Miller, C. O., Skoog, F., von Saltza, M. H., and Strong, F. M. (1955). "Kinetin, a cell division factor from deoxyribonucleic acid". J. Am. Chem. Soc. 77:1392.

Neljubow, D. N. (1901). "Uber die horizontale nutation der stengel von Pisum sativum und einiger anderen". Pflanzen Beitrage und Botanik Zentralblatt 10:128-139.

Paal, A. (1918). "Uber phototropische Reizleitung". Jahrb. Wiss. Bot. 58:406-458.

Radley, M. (1956). "Occurrence of substances similar to gibberellic acid in higher plants". Nature 178:1070-1071.

Raven, P. H., Evert, R. F., and Eichhorn, S. E. (1992). Biology of Plants. New York: Worth. pp. 545-572.

Salisbury, F. B., and Ross, C. W. (1992). Plant Physiology. Belmont, CA: Wadsworth. pp. 357-407, 531-548.

Salkowski, E. (1885). "Uber das verhalten der skatolcarbonsaure im organismus". Zeitschr. Physiol. Chem. 9:23-33.

Sawada, K. (1912). "Disease of agricultural products in Japan". Formosan Agr. Rev. 36:10.

Soding H. (1925). "Zur kenntnis der wuchshormone in der haferkoleoptile". Jahrb. Wiss. Bot. 64:587-603.

Sponsel, V. M. (1995). "Gibberellin biosynthesis and metabolism". Plant Hormones: Physiology, Biochemistry and Molecular Biology. Dordrecht: Kluwer. pp. 66-97.

Stodola, F. H., Raper, K. B., Fennell, D. I., Conway, H. F., Sohns, V. E., Langford, C. T. and Jackson, R. W. (1955). "The microbiological production of gibberellins A and X". Arch. Biochem. Biophys. 54:240-245.

Takahashi, N., Kitamura, H., Kawarada, A., Stea Y., Takai, M., Tamura, S., and Sumiki, Y. (1955). "Isolation of gibberellins and their properties". Bull. Agric. Chem. Soc. Japan 19:267-277.

Takahashi, N., Phinney. B.O., and MacMillan J. (1991). Gibberellins. New York: Springer-Verlag.

Takahashi, N., Seta, Y., Kitamura, H. and Sumiki, Y. (1957). "A new gibberellin, gibberellin A4". Bull. Agric. Chem. Soc. Japan 21:396-398.

van Overbeek, J., Conklin, M. E., and Blakeslee, A. F. (1941). "Factors in coconut milk essential for growth and development of Datura embryos". Science 94:350.

von Sachs, J. (1880). "Stoff und Form der Pflanzenorgane I". Arb. Bot. Inst. Wurzburg 2:452-488.

Walton, D. C., and Li, Y. (1995). "Abscisic acid biosynthesis and metabolism". Plant Hormones: Physiology, Biochemistry and Molecular Biology. Dordrecht: Kluwer. pp. 140-157.

Went, F. W. (1926). "On growth-accelerating substances in the coleoptile of Avena sativa". Proc. Kon. Ned. Akad. Wet. 30:10-19.

Went, F. W. (1928). "Wuchsstoff und Wachstum". Rec. Trav. Bot. Neerland. 24:1-116.

Wolfe, S. L. (1993). Molecular and Cellular Biology. Belmont, CA: Wadsworth. pp. 702-704.

Yabuta, T. (1935). "Biochemistry of the 'bakanae' fungus of rice". Agr. Hort. (Tokyo) 10:17-22.