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Thorns, spines, and prickles

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680: 298: 315: 31: 602: 595:, in particular, are distinct from thorns, spines, and prickles in that they are much smaller (often microscopic) outgrowths of epidermal tissue, and they are less rigid and more hair-like in appearance; they typically consist of just a few cells of the outermost layer of epidermis, whereas prickles may include cortex tissue. Trichomes are often effective defenses against small insect herbivores; thorns, spines, and prickles are usually only effective against larger herbivores like birds and mammals. 469: 451: 656: 411: 433: 239: 620: 570: 227: 215: 487: 537: 2734: 280: 1461: 638: 555: 767:(Rubiaceae), these probably give protection to ants which inhabit chambers within the tuber as they wander over the plant's surface. (Jackson 1986 and references therein). In many respects, the pattern of spine formation is similar to that which occurs in the development of thorns from lateral shoots. (Jackson 1986 and references therein). 257: 731:), there are spiny roots; some authors prefer to term these "root spines" if the length of the root is less than 10x the thickness and "spine roots" if the length is more than 10x the thickness. Adventitious spiny roots have also been described on the trunks of dicotyledonous trees from tropical Africa (e.g. Euphorbiaceae, as in 706:
Lateral roots on these two types of roots, as well as those on the stilt roots on this species, also become spinous. Some authors believe that some of these short spiny laterals have a ventilating function so they are 'pneumorhizae'. Short spiny laterals that may have a ventilating function may also be found on roots of
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grow downwards to a length of 6–12 cm, then stop growing and transform into a spine. The anatomy of crown roots on this species (roots among the bases of the living fronds) also alters during their life. They initially grow upwards and then turn down and finally they, too, become spinous.
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is a term describing plants that bear any sharp structures that deter herbivory. It also can refer to the state of tending to be or become spiny in some sense or degree, as in: "... the division of the African acacias on the basis of spinescent stipules versus non-spinescent stipules..."
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Satterlee, James W.; Alonso, David; Gramazio, Pietro; Jenike, Katharine M.; He, Jia; Arrones, Andrea; Villanueva, Gloria; Plazas, Mariola; Ramakrishnan, Srividya; Benoit, Matthias; Gentile, Iacopo; Hendelman, Anat; Shohat, Hagai; Fitzgerald, Blaine; Robitaille, Gina M. (2024-08-02).
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are limited to defense from physical attacks by herbivores and other animals. In some cases, spines have been shown to shade or insulate the plants that grow them, thereby protecting them from extreme temperatures. For example,
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Ross, J. H. "A conspectus of the African Acacia species." Series: Memoirs of the Botanical Survey of South Africa, No. 44 Botanical Research Institute, Dept. of Agricultural Technical Services, Pretoria,
2564: 1141:"Benefits and Constraints on Plant Defense against Herbivores: Spines Influence the Legitimate and Illegitimate Flower Visitors of Yellow Star Thistle, Centaurea solstitialis L. (Asteraceae)" 920:, being strategically planted below windows or around the entire perimeter of a property. They also have been used to protect crops and livestock against marauding animals. Examples include 2569: 343:, or parts of leaves, such as extensions of leaf veins. Some authors prefer not to distinguish spines from thorns because, like thorns, and unlike prickles, they commonly contain 2702: 314: 394:, singular glochidium) – a particular kind of spine of different origin, which are smaller and deciduous with numerous retrose barbs along its length (as found in 1138:
Agrawal, Anurag A.; Rudgers, Jennifer A.; Botsford, Louis W.; Cutler, David; Gorin, Jessica B.; Lundquist, Carolyn J.; Spitzer, Brian W.; Swann, Alisa L. (2000).
410: 450: 2620: 1469: 679: 601: 2436: 2768: 2763: 380:), all of which are examples of spines developing from a part of a leaf containing the petiole, midrib, or a secondary vein. The plants of the 102:(so that they may or may not be branched, they may or may not have leaves, and they may or may not arise from a bud), spines are derived from 1407: 1378: 1122: 536: 432: 1484: 523:, the positioning of prickles appears to be truly random. If not, then by a phyllotaxis so arcane as to give the appearance of randomness. 1052: 655: 468: 1232: 619: 569: 2626: 1361: 1215: 486: 1521: 191:(2000) found that spines seem to have little effect on specialist pollinators, on which many plants rely in order to reproduce. 1960: 1489: 966: 87: 637: 2441: 211:
prickles) do not have vascular bundles inside, so that they can be removed more easily and cleanly than thorns and spines.
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In common language, the terms are used more or less interchangeably, but in botanical terms, thorns are derived from
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Spinose structures occur in a wide variety of ecologies, and their morphology also varies greatly. They occur as:
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Prickles are comparable to hairs but can be quite coarse (for example, rose prickles). They are extensions of the
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in a mechanical form. For this reason, they are classified as physical or mechanical defenses, as opposed to
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in plants growing in sandy environments that provided inadequate resources for fast regeneration of damage.
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Leaf margins may also have teeth, and if those teeth are sharp, they are called spinose teeth on a spinose
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concluded that plants with these types of prickles have been identified as sharing a common gene family.
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spp. -Iridaceae- respectively). Short root spines cover the tuberous base of the epiphytic ant-plant
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thorn, its leaves, nodes, and emergence from an axillary bud demonstrating its nature as a branch.
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tissue (so that they can be found anywhere on the plant and do not have vascular bundles inside).
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There are also spines derived from roots, like the ones on the trunk of the "Root Spine Palms" (
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Judd, Campbell, Kellogg, Stevens, Donoghue. 2007. "Structural and Biochemical Characters". In:
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Judd, Campbell, Kellogg, Stevens, Donoghue. 2007. "Structural and Biochemical Characters". In:
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Hard, rigid extensions or modifications of leaves, roots, stems or buds with sharp, stiff ends
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are derived from shoots and leaves respectively, and have vascular bundles inside, whereas
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Pointing or spinose processes can broadly be divided by the presence of vascular tissue:
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Other similar structures are spinose teeth, spinose apical processes, and trichomes.
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are particularly well known for their dense covering of spines. Some cacti have also
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Plants bearing thorns, spines, or prickles are often used as a defense against
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in this case; for some authors a kind of prickle), it may be referred to as a
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Dyer, R. Allen, “The Genera of Southern African Flowering Plants”, Vol 2.
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with sharp, stiff ends, and generally serve the same function: physically
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There can be found also spines or spinose structures derived from roots.
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in the Americas and in other countries where they have been introduced,
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It has been proposed that thorny structures may have first evolved as a
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The predominant function of thorns, spines, and prickles is deterring
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There are also spines that function as pneumorhizae on the palm
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Anderson, Edward F., The Cactus Family, Pub: Timber Press 2001
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Plant form: an illustrated guide to flowering plant morphology
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August Weismann, John Arthur Thomson, Margaret R. Thomson,
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Plant Systematics, a phylogenetic approach, third edition
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Hunter, J. A. (1993) "Hunter" Publisher: Buccaneer Books,
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Plant Systematics, a phylogenetic approach, third edition
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a tiny point at the tip of the leaf (mucronate leaves) (
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Identifying and Harvesting Edible and Medicinal Plants
1048: 1046: 1044: 1042: 2676: 2640: 2530: 2403: 2319: 2215: 2165: 1890: 1820: 1763: 1725: 1699: 1635: 1555: 230:Cactus areoles; shoot (yellow), spines (green) and 184:, glochids insulate the apical meristem in winter. 1139: 1028: 1026: 1024: 1022: 1020: 1018: 1016: 1014: 70:), are hard, rigid extensions or modifications of 526:A study published for peer review to the journal 1106: 1104: 1102: 1100: 1098: 1096: 1094: 1092: 1090: 1088: 1086: 1084: 1000: 998: 996: 994: 992: 990: 988: 986: 984: 982: 1515: 1441:, 2nd Edition. John Wiley & Sons. 767 pp. 1423:Simpson, M. G. 2010. "Plant Morphology". In: 1233:"Cross-Section Through the Prickle of a Rose" 1210:(illustrated ed.). Struik. p. 184. 1004:Simpson, M. G. 2010. "Plant Morphology". In: 8: 2621:International Association for Plant Taxonomy 1055:Sonoran Desert Plants, an Ecological Atlas. 1722: 1522: 1508: 1500: 1291: 1117:. Springer Science & Business Media. 1114:New Root Formation in Plants and Cuttings 2565:International Code of Nomenclature (ICN) 1208:How to Identify Trees in Southern Africa 1193:. Oxford University Press, Oxford, U.K. 978: 597: 532: 406: 325:showing characteristic branched thorns. 275: 1185: 1183: 1075:How to identify trees in South Africa. 195:Definitions and technical distinctions 1427:. Elsevier Academic Press. Chapter 9. 1008:. Elsevier Academic Press. Chapter 9. 936:in the prairie states of the US, and 7: 456:Areoles and spines of the tree-like 839:stipules converted to spines (many 886:Some thorns are hollow and act as 830:leaves fully converted to spines ( 350:Spines are variously described as 273:. They may be simple or branched. 88:defending plants against herbivory 25: 2570:ICN for Cultivated Plants (ICNCP) 1448:. Timber Press, Portland. 423 pp. 180:in summer, and in members of the 114:), and prickles are derived from 2733: 2732: 1490:The New Student's Reference Work 1459: 870:urticating (i.e. stinging) hairs 683:"Root spines" on the trunk of a 654: 636: 618: 600: 568: 553: 535: 485: 467: 449: 431: 409: 313: 296: 278: 269:Thorns are modified branches or 252:A spinose tooth in a leaf margin 1425:Plant Systematics, 2nd. edition 1231:Sengbusch, Peter (2003-07-31). 1006:Plant Systematics, 2nd. edition 967:Plant defense against herbivory 542:Raised prickles on the stem of 234:(also spines, green and little) 167:Not all functions of spines or 1: 2769:Hazards of outdoor recreation 2764:Biological defense mechanisms 1111:Jackson, M. B. (2012-12-06). 2611:History of plant systematics 2198:Thorns, spines, and prickles 1057:University of Arizona Press. 878:finely barbed spines called 791:sharpened branches (e.g. in 1146:The Southwestern Naturalist 18:Thorns, spines and prickles 2785: 2427:Alternation of generations 1475:Collier's New Encyclopedia 699:spp.). The trunk roots of 643:Stiff, sharp trichomes in 625:Spinose apical process in 575:Prickles on the leaves of 2728: 2577:Cultivated plant taxonomy 2540:Biological classification 1537: 1446:Tropical Flowering Plants 2437:Evolutionary development 1073:Van Wyk, Van Wyk. 2007. 459:Rhodocactus grandifolius 260:A spinose apical process 68:spinose apical processes 2088:Hypanthium (Floral cup) 1276:10.1126/science.ado1663 1206:Van Wyk, Braam (2007). 1195:preview in google books 783:Morphological variation 607:Spinose leaf margin in 495:Mammillaria balsasoides 133:(more correctly called 2703:by author abbreviation 2627:Plant taxonomy systems 2545:Botanical nomenclature 1389:Felson, Marcus (2006) 812:spiky inflorescences ( 690: 545:Caesalpinia decapetala 441:Vachellia xanthophloea 422:develop from the leaf 261: 253: 245: 235: 223: 127:pungent apical process 39: 2710:Botanical expeditions 1464:Texts on Wikisource: 752:Dioscorea prehensilis 729:Acanthorhiza aculeata 682: 661:Stinging trichome in 305:Gymnosporia buxifolia 259: 251: 241: 229: 217: 33: 2442:Evolutionary history 2432:Double fertilization 2284:Cellular respiration 1444:Llamas, K. A. 2003. 1334:The Evolution Theory 1053:Turner et al. 2005, 899:Crataegus crus-galli 755:-Dioscoreaceae- and 419:Fouquieria splendens 335:Spines are modified 285:Smooth, featureless 1661:Non-vascular plants 848:prickles on stems ( 764:Myrmecodia tuberosa 702:Cryosophila guagara 222:and arise from buds 2166:Surface structures 1961:Flower development 924:hedges in Europe, 905:Carissa macrocarpa 893:Crataegus monogyna 890:; others (e.g. in 740:Bridelia micrantha 709:Iriartea exorrhiza 691: 438:Stipule spines on 262: 254: 246: 242:(A) Thorn or spine 236: 224: 62:(sometimes called 60:spinose structures 40: 2746: 2745: 2385:Herbaceous plants 2211: 2210: 1408:978-1-56849-109-7 1379:978-0-88192-498-5 1189:Bell, A.D. 1997. 1124:978-94-009-4358-2 777:defense mechanism 734:Macaranga barteri 322:Carissa bispinosa 201:thorns and spines 176:spines shade the 58:, and in general 16:(Redirected from 2776: 2759:Plant morphology 2736: 2735: 2715:Individual trees 2390:Secondary growth 2361:Succulent plants 2349:Prostrate shrubs 2232:Apical dominance 2217:Plant physiology 2178:Epicuticular wax 1723: 1716: 1707:Plant morphology 1524: 1517: 1510: 1501: 1494: 1479: 1463: 1410: 1400: 1394: 1391:Crime and Nature 1387: 1381: 1371: 1365: 1354: 1348: 1343: 1337: 1330: 1324: 1314: 1308: 1304: 1298: 1297: 1295: 1254: 1248: 1247: 1245: 1244: 1235:. 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947: 944: 913: 910: 888:myrmecodomatia 884: 883: 876: 873: 867: 863:Ceiba speciosa 846: 837: 828: 819: 810: 784: 781: 772: 769: 670: 669: 660: 653: 651: 646:Galium aparine 642: 635: 633: 624: 617: 615: 606: 599: 588: 585: 584: 583: 578:Solanum viarum 574: 567: 565: 559: 552: 550: 541: 534: 504: 501: 500: 499: 491: 484: 482: 473: 466: 464: 455: 448: 446: 437: 430: 428: 416:The spines of 415: 408: 362:leaflet spines 332: 329: 328: 327: 319: 312: 310: 302: 295: 293: 284: 277: 266: 263: 220:modified stems 196: 193: 174:saguaro cactus 153: 150: 95: 92: 34:Prickles on a 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 2781: 2770: 2767: 2765: 2762: 2760: 2757: 2756: 2754: 2739: 2731: 2730: 2727: 2721: 2718: 2716: 2713: 2711: 2708: 2704: 2701: 2700: 2699: 2696: 2694: 2691: 2690: 2688: 2682: 2679: 2678: 2675: 2669: 2668:Phytochemical 2666: 2664: 2661: 2659: 2656: 2654: 2651: 2649: 2646: 2645: 2643: 2639: 2633: 2630: 2628: 2625: 2622: 2619: 2617: 2614: 2612: 2609: 2603: 2600: 2598: 2595: 2593: 2590: 2589: 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2168: 2164: 2158: 2155: 2153: 2150: 2148: 2145: 2143: 2140: 2136: 2133: 2131: 2128: 2126: 2123: 2122: 2121: 2118: 2114: 2111: 2109: 2106: 2104: 2101: 2099: 2096: 2095: 2094: 2093:Inflorescence 2091: 2089: 2086: 2080: 2077: 2075: 2072: 2068: 2065: 2063: 2060: 2059: 2058: 2055: 2054: 2053: 2050: 2049: 2048: 2045: 2043: 2040: 2038: 2035: 2029: 2026: 2024: 2021: 2020: 2019: 2016: 2014: 2011: 2009: 2006: 2004: 2001: 1999: 1996: 1994: 1991: 1990: 1989: 1986: 1982: 1979: 1977: 1974: 1972: 1969: 1967: 1964: 1962: 1959: 1957: 1954: 1953: 1952: 1949: 1945: 1942: 1940: 1937: 1933: 1930: 1928: 1925: 1924: 1923: 1920: 1918: 1915: 1914: 1913: 1910: 1908: 1905: 1903: 1900: 1899: 1897: 1893: 1889: 1881: 1878: 1876: 1873: 1869: 1866: 1864: 1861: 1860: 1859: 1856: 1854: 1851: 1850: 1849: 1846: 1844: 1841: 1839: 1836: 1834: 1831: 1829: 1826: 1825: 1823: 1819: 1813: 1810: 1806: 1803: 1802: 1801: 1798: 1796: 1793: 1791: 1788: 1784: 1781: 1780: 1779: 1778:Ground tissue 1776: 1774: 1771: 1770: 1768: 1766: 1762: 1756: 1753: 1751: 1748: 1746: 1743: 1741: 1738: 1736: 1733: 1732: 1730: 1728: 1724: 1721: 1714: 1708: 1705: 1704: 1702: 1701:Plant anatomy 1698: 1692: 1689: 1687: 1684: 1682: 1679: 1677: 1674: 1672: 1669: 1667: 1664: 1662: 1659: 1657: 1654: 1652: 1649: 1647: 1644: 1643: 1641: 1638: 1634: 1628: 1625: 1623: 1622:Plant ecology 1620: 1618: 1617:Plant anatomy 1615: 1611: 1608: 1607: 1606: 1603: 1601: 1598: 1596: 1593: 1591: 1588: 1586: 1583: 1581: 1578: 1576: 1573: 1571: 1568: 1566: 1565:Archaeobotany 1563: 1562: 1560: 1558: 1554: 1548: 1545: 1543: 1540: 1539: 1536: 1532: 1525: 1520: 1518: 1513: 1511: 1506: 1505: 1502: 1492: 1491: 1486: 1481: 1477: 1476: 1471: 1470:Thorn (plant) 1466: 1465: 1462: 1458: 1457: 1453: 1447: 1443: 1440: 1439:Plant Anatomy 1436: 1433: 1429: 1426: 1422: 1421: 1416: 1409: 1405: 1399: 1396: 1392: 1386: 1383: 1380: 1376: 1370: 1367: 1363: 1362:0-621-02863-0 1359: 1353: 1350: 1347: 1342: 1339: 1335: 1329: 1326: 1322: 1318: 1313: 1310: 1303: 1300: 1294: 1289: 1285: 1281: 1277: 1273: 1269: 1265: 1261: 1253: 1250: 1239:on 2008-04-30 1238: 1234: 1227: 1224: 1219: 1217:9781770072404 1213: 1209: 1202: 1199: 1196: 1192: 1186: 1184: 1180: 1175: 1171: 1167: 1163: 1159: 1155: 1151: 1147: 1142: 1134: 1131: 1126: 1120: 1116: 1115: 1107: 1105: 1103: 1101: 1099: 1097: 1095: 1093: 1091: 1089: 1087: 1085: 1081: 1078: 1076: 1069: 1067: 1065: 1061: 1058: 1056: 1049: 1047: 1045: 1043: 1039: 1035: 1029: 1027: 1025: 1023: 1021: 1019: 1017: 1015: 1011: 1007: 1001: 999: 997: 995: 993: 991: 989: 987: 985: 983: 979: 972: 968: 965: 963: 960: 958: 955: 953: 950: 949: 945: 943: 941: 940: 935: 931: 927: 923: 919: 911: 909: 907: 906: 901: 900: 895: 894: 889: 881: 877: 875:bristles, and 874: 871: 868: 865: 864: 859: 858: 853: 852: 847: 844: 843: 838: 835: 834: 829: 826: 825: 820: 817: 816: 811: 808: 807: 802: 801: 796: 795: 790: 789: 788: 782: 780: 778: 770: 768: 766: 765: 760: 759: 754: 753: 748: 747: 742: 741: 736: 735: 730: 726: 725: 720: 719: 713: 711: 710: 704: 703: 698: 697: 688: 687: 681: 677: 674: 666: 665: 664:Urtica dioica 657: 652: 648: 647: 639: 634: 630: 629: 621: 616: 612: 611: 603: 598: 596: 594: 586: 580: 579: 571: 566: 562: 556: 551: 547: 546: 538: 533: 531: 529: 524: 522: 518: 514: 510: 502: 497: 496: 488: 483: 480: 478: 470: 465: 461: 460: 452: 447: 443: 442: 434: 429: 425: 421: 420: 412: 407: 405: 403: 402: 397: 393: 389: 388: 383: 382:cactus family 379: 378: 373: 369: 368: 363: 359: 358: 353: 348: 346: 342: 338: 330: 324: 323: 316: 311: 307: 306: 299: 294: 290: 289: 281: 276: 274: 272: 264: 258: 250: 240: 233: 228: 221: 216: 212: 210: 206: 202: 194: 192: 190: 185: 183: 179: 175: 170: 165: 163: 159: 151: 149: 146: 144: 140: 136: 132: 128: 124: 119: 117: 113: 109: 105: 101: 93: 91: 89: 85: 81: 77: 73: 69: 65: 64:spinose teeth 61: 57: 53: 49: 45: 37: 32: 19: 2663:Horticulture 2653:Floriculture 2555:Correct name 2405:Reproduction 2395:Woody plants 2320:Plant growth 2279:Gas Exchange 2264:Phytomelanin 2197: 2142:Plant embryo 1892:Reproductive 1740:Phragmoplast 1488: 1473: 1445: 1438: 1434:. Chapter 4. 1431: 1424: 1417:Bibliography 1398: 1390: 1385: 1369: 1352: 1346:Bihrmann.com 1341: 1333: 1328: 1320: 1312: 1302: 1267: 1263: 1252: 1241:. Retrieved 1237:the original 1226: 1207: 1201: 1190: 1149: 1145: 1133: 1113: 1074: 1054: 1036:. Chapter 4. 1033: 1005: 937: 934:Osage orange 915: 903: 897: 891: 885: 861: 855: 849: 840: 831: 822: 813: 804: 798: 792: 786: 774: 762: 756: 750: 746:B. pubescens 744: 738: 732: 728: 722: 716: 714: 707: 700: 694: 692: 684: 672: 671: 662: 644: 626: 608: 590: 576: 543: 525: 506: 493: 475: 457: 439: 417: 399: 391: 385: 375: 371: 365: 361: 355: 351: 349: 334: 320: 303: 286: 268: 204: 200: 198: 188: 186: 182:Opuntioideae 166: 155: 147: 142: 138: 126: 120: 97: 67: 63: 59: 55: 51: 47: 41: 2479:Pollen tube 2474:Pollinators 2464:Pollination 2459:Germination 2274:Respiration 2257:Chlorophyll 2103:Pedicellate 2037:Gametophyte 1956:Aestivation 1907:Antheridium 1902:Archegonium 1750:Plasmodesma 1727:Plant cells 1590:Paleobotany 1585:Ethnobotany 1570:Astrobotany 1317:Steve Brill 942:in Africa. 939:Sansevieria 824:Sansevieria 696:Cryosophila 686:Cryosophila 628:Sansevieria 521:phyllotaxis 479:balsasoides 477:Mammillaria 244:(B) Prickle 218:Thorns are 123:leaf margin 94:Description 2753:Categories 2501:Microspore 2491:Sporangium 2469:Artificial 2157:Sporophyte 2152:Sporophyll 2147:Receptacle 2042:Gynandrium 1912:Androecium 1821:Vegetative 1691:Angiosperm 1686:Gymnosperm 1580:Dendrology 1243:2009-04-27 1152:(1): 1–5. 973:References 912:Human uses 727:(formerly 673:Spinescent 474:Spines of 357:Fouquieria 164:defenses. 36:blackberry 2698:Botanists 2616:Herbarium 2513:Megaspore 2411:Evolution 2354:Subshrubs 2322:and habit 2247:Nutrition 2242:Cellulose 2237:Bulk flow 2220:Materials 2183:Epidermis 2047:Gynoecium 2028:Endosperm 2023:Dispersal 1939:Staminode 1875:Sessility 1863:Cataphyll 1783:Mesophyll 1735:Cell wall 1676:Lycophyte 1656:Bryophyte 1610:Geobotany 1595:Phycology 1393:, p. 288. 1284:0036-8075 1166:0038-4909 930:ocotillos 857:Erythrina 842:Vachellia 815:Tylecodon 806:Crataegus 771:Evolution 593:Trichomes 563:prickles. 513:epidermis 392:glochidia 377:Euphorbia 158:herbivory 131:trichomes 116:epidermis 2738:Category 2658:Forestry 2648:Agronomy 2641:Practice 2592:Cultivar 2587:Cultigen 2447:timeline 2339:Rosettes 2227:Aleurone 2203:Trichome 2120:Perianth 1932:Filament 1790:Meristem 1713:glossary 1575:Bryology 1293:11305333 1270:(6708). 962:Trichome 946:See also 922:hawthorn 918:burglary 880:glochids 689:species. 503:Prickles 424:petioles 387:glochids 341:stipules 232:glochids 205:prickles 187:Agrawal 169:glochids 162:chemical 152:Function 135:bristles 56:prickles 2416:Ecology 2173:Cuticle 2003:Capsule 1993:Anatomy 1944:Tapetum 1868:Petiole 1843:Rhizome 1838:Rhizoid 1765:Tissues 1755:Vacuole 1745:Plastid 1547:Outline 1542:History 1493:. 1914. 1478:. 1921. 1264:Science 1174:3672545 1077:Struik. 957:Glochid 833:Opuntia 794:Carissa 528:Science 401:Opuntia 396:areoles 374:(as in 367:Phoenix 364:(as in 354:(as in 112:stipule 108:petiole 2720:Plants 2623:(IAPT) 2376:Lianas 2344:Shrubs 2296:Starch 2188:Nectar 2108:Raceme 2074:Stigma 2062:Locule 2052:Carpel 2013:Pyrena 1951:Flower 1927:Anther 1922:Stamen 1917:Pollen 1639:groups 1531:Botany 1406:  1377:  1364:, 1976 1360:  1290:  1282:  1214:  1172:  1164:  1121:  952:Areole 926:agaves 800:Citrus 758:Moraea 509:cortex 370:), or 337:leaves 331:Spines 291:thorn. 288:Citrus 265:Thorns 207:(like 189:et al. 104:leaves 100:shoots 72:leaves 54:, and 52:spines 48:thorns 38:branch 2680:Lists 2597:Group 2520:Spore 2454:Flora 2371:Vines 2366:Trees 2329:Habit 2301:Sugar 2193:Stoma 2135:Sepal 2130:Petal 2125:Tepal 2113:Umbel 2098:Bract 2079:Style 2067:Ovule 2057:Ovary 1998:Berry 1988:Fruit 1981:Whorl 1848:Shoot 1646:Algae 1637:Plant 1485:Thorn 1170:JSTOR 721:. In 517:Roses 271:stems 110:or a 82:, or 80:stems 76:roots 2602:Grex 2484:Self 2018:Seed 1880:Stem 1858:Leaf 1833:Root 1828:Bulb 1812:Wood 1773:Cork 1671:Fern 1404:ISBN 1375:ISBN 1358:ISBN 1307:1979 1280:ISSN 1212:ISBN 1162:ISSN 1119:ISBN 902:and 860:and 851:Rosa 743:and 561:Rose 511:and 390:(or 209:rose 84:buds 2291:Sap 2008:Nut 1853:Bud 1487:". 1472:". 1288:PMC 1272:doi 1268:385 1154:doi 928:or 908:). 404:). 398:of 360:), 66:or 42:In 2755:: 1286:. 1278:. 1266:. 1262:. 1182:^ 1168:. 1160:. 1150:45 1148:. 1144:. 1083:^ 1063:^ 1041:^ 1013:^ 981:^ 866:), 854:, 845:), 836:), 827:), 818:), 809:), 803:, 797:, 737:, 712:. 347:. 339:, 145:. 90:. 78:, 74:, 50:, 46:, 1715:) 1711:( 1523:e 1516:t 1509:v 1483:" 1468:" 1296:. 1274:: 1246:. 1220:. 1176:. 1156:: 1127:. 882:. 872:, 667:. 649:. 631:. 613:. 581:. 548:. 462:. 444:. 426:. 20:)

Index

Thorns, spines and prickles

blackberry
plant morphology
leaves
roots
stems
buds
defending plants against herbivory
shoots
leaves
petiole
stipule
epidermis
leaf margin
trichomes
bristles
herbivory
chemical
glochids
saguaro cactus
apical meristem
Opuntioideae
rose

modified stems

glochids

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