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Uterotonic Plants and Their Bioactive Constituents

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Uterotonic Plants and Their Bioactive Constituents Reviews 207 Uterotonic Plants and their Bioactive Constituents Authors Christian W. Gruber1, Margaret OʼBrien2 Affiliations 1 Department of Pharmacognosy, University of Vienna, Vienna, Austria 2 National Centre for Biomedical and Engineering Science, Orbsen Building, National University of Ireland Galway, Galway, Ireland Key words Abstract their mechanisms, primarily focusing on utero- l" womenʼs health ! tonic active circular plant peptides called cycloti- l" gynaecology Abnormalities in the process of uterine muscle des. Finally we will comment on the discovery of l" labour contractility during pregnancy and birth can have novel cyclotide-producing plant species and the l" myometrial smooth muscle major clinical implications, including preterm la- possibility for the development of novel plant-de- l" oxytocin l" cyclotides bour, which is the single largest cause of maternal rived uterotonic and tocolytic drugs. l" plants and prenatal mortality in the Western world and a major contributor to childhood developmental problems. In contrast, induction of labour may Abbreviations be necessary in certain conditions. Currently used ! interventional therapies to suppress (tocolytic AC: adenylyl cyclase agents) or to induce (uterotonic agents) uterine CCK: cyclic cystine knot contractions lack potency and/or selectivity and COX-2: cyclooxygenase-2 can have harmful side effects for mother and CRH: corticotropin-releasing hormone baby. Natureʼs diversity has always been, and still GPCR: G-protein coupled receptor is, one of the biggest resources of therapeutic lead IL: interleukin compounds. Many natural products exhibit bio- IP3: D-myoinositol 1,4,5-triphosphate logical activity against unrelated targets, thus MLC: myosin light chain providing researchers with starting points for NF: nuclear factor drug development. In this review we will provide OT: oxytocin an overview of uterine muscle physiology, de- PL: phospholipase scribe currently available biological screening R: receptor procedures for testing of uterotonic plant com- SFE: supercritical fluid extraction received June 1, 2010 pounds and will summarise traditionally-used SR: sarcoplasmic reticulum revised July 28, 2010 uterotonic plants, their active components and TNF: tumour necrosis factor accepted August 5, 2010 Bibliography DOI http://dx.doi.org/ Introduction volve the modulation of uterine contractions at 10.1055/s-0030-1250317 ! labour, resulting in either the stimulation (“utero- Published online September 15, Traditional medicine relies on the use of certain tonic”) or inhibition (“tocolytic”) of myometrial 2010 herbal plants and other remedies for beneficial ef- muscle contractions. In this review we will pro- Planta Med 2011; 77: 207–220 © Georg Thieme Verlag KG fects during pregnancy, to induce labour, in the vide an overview of uterotonic herbs and their ac- Stuttgart · New York · removal of retained placenta and management of tive constituents. This includes traditionally-used ISSN 0032‑0943 post-partum bleeding. However, some of these as well as recently identified plants or isolated Correspondence medicines have harmful side effects and when plant compounds, which have reportedly shown Dr. Christian W. Gruber taken in large quantities can lead to the death of uterine smooth muscle-stimulating activities. Department of Pharmacognosy the unborn baby and/or uterine rupture, and oth- Furthermore we will give a brief outline of avail- University of Vienna Althanstrasse 14 er longer term effects on the mother or baby. able uterotonic assay systems and provide a list 1090 Vienna Most often the biological effects elicited by these of uterotonic plants, their bioactive ingredients Austria remedies are due to biomolecules (small chemi- and their mechanism of action on the uterus. Our Phone: + 431427755262 Fax: + 431 42779552 cals, peptides or proteins) that primarily act on focus will be on the description of recently identi- [email protected] the uterus. The nature of these actions may in- fied cyclotide peptides, mainly found in plants Gruber CW, OʼBrien M. Uterotonic Plants and… Planta Med 2011; 77: 207–220 208 Reviews from the Rubiaceae and Violaceae families, which have shown to be uterotonic agents. We will begin this article with a thorough description of uterine physiology and molecular details of uter- ine smooth muscle contractility. Uterine Muscle Physiology ! Biology of the uterus The uterus is the central organ of reproduction. It is a thick, pear- shaped, muscular organ approximately, 7 cm long, and 4–5cm wide at its widest point. It is divided functionally and morpho- logically into three sections, namely the cervix, the isthmus and the main body of the uterus (corpus uteri) [1]. The myometrium is the middle muscular layer that makes up the major proportion of the uterine body. Myometrial smooth muscle is arranged in undefined layers and contractile forces can occur in any direction enabling the uterus to assume virtually any shape. Fig. 1 Uterine contraction signalling pathways. Uterotonic pathways (red Through growth and stretch during pregnancy, the myometrium arrows) – Myometrial contraction and relaxation result from the phospho- provides the protective environment for the developing foetus. rylation or dephosphorylation of myosin light chains (MLC), respectively. Then with the onset of labour it contracts rhythmically to expel Phosphorylation, by the enzyme myosin light chain kinase (MLCK), in the presence of adenosine triphosphate (ATP), is regulated by intracellular cal- the foetus and placenta. Smooth muscle fibres are composed of 2+ cium concentrations ([Ca ]i), in conjunction with the intermediate protein spindle-shaped cells, each with one centrally located nucleus. Typ- calmodulin (CaM), which together form the calcium-calmodulin (Ca-CaM) ically, they have a diameter of 2–10 µm and a length of several complex. Calcium channels (voltage and agonist operated channels; VOC/ hundred µm [2]. Smooth muscle cells are embedded in an extra- AOC), membrane endothelin (ET) receptors (ET1,ET2,ET3), passive entry, cellular matrix composed principally of collagen fibres, which fa- membrane prostaglandin (PG) receptors (E2,F2α) and stretch, all facilitate 2+ ↑ 2+ cilitate the transmission of contractile forcesgenerated by individ- an increase in intracellular Ca concentration ( [Ca ]i) and result in ual cells. They are organised into sheets of closely opposed fibres, smooth muscle contraction. Agonist-mediated activation of membrane acetylcholine (ACh) and oxytocin (OT) receptors stimulate the production oriented at right angles to each other. These sheets form two dis- of the second messenger D-myoinositol 1,4,5-triphosphate (IP ), the latter “ ” 3 tinct layers, the longitudinal layer , which consists of a network through the action of the enzyme phospholipase C (coupled to the OTR by of bundles of smooth muscle cells generally oriented in the long the G-protein [Gq]) on the plasma membrane constituent phosphatidyl- 2+ axis of the organ, and the “circular layer”, in which the fibres are inositol 4,5-bisphosphate (PIP2). IP3 releases Ca from the sarcoplasmic 2+ arranged concentrically around the longitudinal axis of the organ reticulum (SR) thus increasing [Ca ]i and resulting in cell contraction. A by- [3]. Contraction of the longitudinal layer causes the organ to dilate product of IP3 synthesis, the second messenger diacylglycerol (DAG) might and shorten, whereas contraction of the circular layer causes the promote cell contraction via intracellular PG synthesis from arachidonic ac- id (AA) by cyclooxygenase (COX) enzymes. The steroid hormone oestrogen organ to elongate; thus alternating contraction and relaxation of promotes cellular contractility by upregulating COX enzymes, particularly these layers enables the uterus to expel its contents at birth [2]. the COX-2 isoform. The active isoform of myosin phosphatase (MP) de- phosphorylates MLC, promoting cell relaxation. Receptor-agonist binding Uterine muscle contractility and the formation or upregulation of intracellular RhoA or Rho kinase could The onset of labour is facilitated by phasic myometrial contrac- result in a shift in the equilibrium of intracellular MP in the direction of the tions that are driven by the development of action potentials inactive isoform, resulting in enhanced cell contraction, i.e., calcium sensi- α α across the plasma membrane, resulting from a transient increase tisation. Agonist binding of the 1 adrenergic receptor ( 1ADR) stimulates inhibitory G-proteins (G ), which inactivate the adenylyl cyclase (AC) medi- in the cytosolic free Ca2+ concentration [4]. In this case (calcium- i ated production of cAMP from ATP. cAMP results in cell relaxation in many dependent smooth muscle contraction), calcium is released from 2+ ways, including inhibition of MLCK and the efflux of [Ca ]i through sodium/ the sarcoplasmic reticulum (SR) and from extracellular stores calcium (Na+/Ca2+) exchanger channels. Chloride (Cl−) channels, which through voltage-gated calcium channels. Therefore smooth might be activated by OT, exert their uterotonic effect by depolarisation of muscle contractility by different agonists or by electrical depolar- the smooth muscle cell membrane. Uterorelaxant pathways are indicated by 2+ 2+ blue arrows. Figure was adapted with modifications from [154]. isation results in a rapid increase in [Ca ]i [5]. Ca binds to four binding sites of calmodulin causing a conformational change al- lowing the calmodulin-calcium complex to interact with inactive
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