teres : © Distribution: limited. wwsco/ss/az/ia5,S Rey / Provisional reproduction PARIS, 31 March “without figures or bibliogrephy Original: Engi UNITED NATIONS EDUCATIONAL, SCIENTIFIC AND CULTURAL ORGANIZATION Major Project _on Scientific Research:on Arid Landg ADAPTATION 10 DROUGHT. : XEROPHYTISM we He R. OPPENHEIMER Faculty of Agrictilture, The Hebrew University, : Rebovot, Israel DEFINITION Generally speaking, plait life presupposes ample moisture. It attains highest development in rainy climates and is handicapped by prolonged reinless periods pecultar tO semi-arid and arid countries. Such conditions threaten plants with desiccation or, to say the least, they impair the aecumlation of ~ assimilated food.end growth, They often: bring about the shedding of flowers and of fruit or the shrivelling of grains and seeds. c Naturel selection and evolution have produced plant, types highly fit to survive under'the hard conditions prevailifig in semi-arid.end.arid countries. Rot only can these plants withstand seiditorms ool atrogons. | Doteptnts mane ‘these plants . = dry, flaccid, and phyton = plant). | The copies of thelr adaptive properties ‘ta dalled Xerophytion. Originated by Schouw (194) so. 1802, const, not easy to define in satisfactory manner. In contrast with “hygrophytes" living in vet habitats, . xerophyte could be any plant living fn dry localities, izrespective either of its activity end rest seasons or of vhether ‘it possesses or lacks the morphological 6 ‘understandNS/AZ/415 - page 2 Even perennials with water-storing subterrenean organs do not necessarily possess obvious adaptive characters, called "xeromorphoses", in their aerial organs. Since their period of vegetative activity is often extrenely limited, ‘they have been called "seasonal ephemerials". ‘The above mentioned difficulties found an early expression in the classifica- tion of plants of dry regions into drought “escaping”, “evading” and “enduring” types by Kearney and Shantz (87).' Their classification was based on ideas first formlated by teh Hyck at the Dry Farming Congress at Billings (Montana) in 1910. ‘The first group includes plants of a short annual growth cycle coinciding with the desert ephemeres. The plants in the second group "evade" fatal effects by restriction of water expense or by extensive but not necessarily deep root systems, or by both. In this group we find numerous adaptive morphological characters. Finally the drought "enduring" plants are able to maintain life activities even when practically no absorption of water by the roats can take place. These include cacti, African euphorbias and other succulents, leafless non-succulent desert shrubs, certain broans and hard-leaved perennial grasses living through the ry season in a more’or less dormant condition, This group includes types with most pronounced adaptive characteristics for the preservation and the economical use of vater. While many earlier ecologists defined xerophytes according either to their topographical, morphological or anatomical adaptive properties, Lene (103) speaks af "physiological xerophytes". This definition implies that this ecological group is well adapted to drought conditions without possessing Obvious morphological characters’ suggesting such adaptability. Genkel (50) recently proposed a definition cambining topogrephical and biological principles. His definition reads: "We call xerophytes plants of dry habitats able to adapt. then- selves successfully to the unfavoureble influence’ of atmospheric and edaphic @rought during their individual develorment thanks to special anatomical- Physiological properties". This definition is good enough though we should prefer to add alternatively “morphological” to the anatanical or physiological properties thus including geopliytes, thorny shrubs, brooms; ete., the adsptation of which is obvious to the naked eye. Schimper (195) 1s often cited for defining xerophytes as plants living under conditions of vater shortage and endowed with the capacity to restrict transpire~ tion, This definition was found inadequate when Maximov (122) shoved that certain xerophytes spend more water than "nesophytes" (plants of medium vet localities) at least vhen transpiration is related to the surface of their leaves. Despite Meximov's criticiom, 1t camnot be denied that arrangements for the restriction of water evaporation from the aerial organs are obvious and an essential property of many plants groving in dry surroundings and of the majority of extrene xerophytes Little studied by him (Zchary; 260). Instead Maximov tried another definition describing xerophytes plants capable of enduring prolonged dehydration often accompanied by permanent wilting. This definition enbraces only the érought- eveding plants of Kearney and Shantz, but is not adequate for the escaping and many enduring types which quite often will show no wilting at all. Another dravback for @ definition based on permanent.wilting is the difficulty of defining wilting scientifically, Obnoxious dehydration is not always matched by a flaccid condition of the plant and it seems preferable to emphasize ition - as Maximov did in later publications - and often overheating, the ws/039.41.NS/AZ/415 .- page 5 nommal development of physiological activities. Since both Schimper's and Maximov's definitions are too narrow to cover all the relevant plant types, ecologists often speak of a xerophyte in the sense of the former or of the latter if- the organism does not combine both properties: low trenspiration at least in tines of restricted water supply and endurance of far-reaching dehydration of its organs. Modern research bas shown that, in contrast to earlier assumptions xerophytes are plants of widely varying character. Stocker (215) stressed the fact that “everywhere we meet with extensive series of different physiological: types differing often much more from each other in the same habitat than corresponding types of widely differing habitats". ‘The came diversity had even earlier been stressed by Keller (88) who compared the lump of different adaptive characteristics amaseed by ecologists together, to parts of different. machines thrown together on one heap. “The "thrilling" diversity of adaptational means belittled by Walter (2h1) was on the other hand stressed by Shreve (205), one of the greatest authori- ties on plant-life in deserts. He insisted on the fact that physiological adaptations are not all matched by structural ones and that, accordingly, each plant type must be analysed physiologically and structurally throughout its evelopment cycle before the complex of its adsptations can be defined. The diversity of structural and logical types in xerophytiam is also stressed ‘by Sveshnikove and Zalensky (221). -If. structural peculiarities, such as thick and cutinized epidermal valls, sclerenchymatic hypodermal tissues, sunken stomta, hard and thickish leaves of reduced size, etc. are present, the plant is called "zeromorphous", a term more definite and adequate than the earlier one “xerophilous" (= fond of drought) which was based on a misinterpretation of the. plant's preferences. Indeed, few plants.of dry habitats do not prefer moist to ary conditions. “Already in 190k Spalding has shown that the creosote bush of the Arizona desert prefers moist habitats to dry ones.and will even plunge its roots into water if offered the opportunity (210). The Freoent writer also observed in Israel the analogous behaviour of the desert palm We filsrera escaped from cultivation (in Wadi Dustre, near the Cribaders' Gastle of Renita). Even cacti observed by ecologists in their native habitats like high moisture in hot weather, (Cannon; 31) a fact well known also to horticulturs ists grdving then commercially. If true xerophytes are not all xeromorphous, conversely xeromorphous plants are not all xerophytes. Thus sclerophyllous (hard-leaved) plants widespread in countries vith’a mediterranean climate are also found in wet tropical and even beyond the arctic cycle, e.g. Rhododendron cum in Lap! needle-leaved shrubs are widespread in oceanic regions, arctic coast of the Kola peninsular. This suggests that seme structural property my serve more than one purpose ani that physiological conjectures based on structural peculfarities - cherished by, ecologists of the past = mst always be corroborated or disproved by experimental methods.(Stocker, 21! De SURVEY OF ADAPTIVE CHARACTERS ‘The present knowledge of the siorphological and anatomical characters of plants growing in dry countries leads back to the fundamental contributions af G, Volkens who was a pupil of 8. Schwendener, originator of physiological principles in plant anatomy. Volkens wrote his famous monograph on the plants W8/039.42.WS/AZ/415 - page 4 of the Egyptian desert (235), but also investigated by experimental methods (232), the influence of axy conditions on European plants, such as Veronica and Asperula, ‘The fundamental principles he established are: (1) ‘he reduction of the amber of size of leaves, their curvature or folding. The reduction of the mmber of leaves con eventually lead to their complete suppression. (2) The principle of water economy in times of drought can also be achieved by the Of the protective tissue. Thickened epidermis valls, their impreg- nation vith cutin as. well as their covering vith waxy bloam serve the. purpose. The resistence to water vapour passing through the stomatais increased by the engthening.of the. way to the surface; we find then elther sunken below the epidermis level, ox concentrated in grooves, as in the desert broom Retama roetam or in cavities, as in the oleander bush. Volkens (232; p.4) insists-on the important ‘reduction in transpiration rete vhich mst result if water supply lags ‘behind transpiretion and an external layer of the cell menbranes becomes msaturated with water. He insists on the fact that this layer must be less permeable to weter than the rest of the membrane, due to "the hindrance met by the vapour molecules thanks to the closing together of the micelles "composing the shrinking cell wells. These highly important and fundamental passages. remained umoticed ty, jater euthore, such ac Renner (185) carrying one respectful controversy against Volkeas and Livingston end Brown vho are credited with the discovery of the Principle of “incipient drying” in plant transpiration. (3) The epidermis can be reinforced by hypodermous layers of mechanical tissue, such as plate collenchyma or bast fibres, or covered by dead hairs reducing. radiation and the velocity of air movements. The increasing proportion of mechanical tissue was recognized as a general principle in the structure of plants growing in dry surroundings. (4) ‘these plants also evince a closed inner structure with reduced intercellular @paces. The differentiation of the mesophyll (internal leaf tissue) into crowded and elongated palisades and spongy parenchyma - the former occurring sometimes in several layers under dry conditions, instead of one under wet conditions -.was shown by Volkens to be not an effect of light and not of drought. Nevertheless, it could be considered as an effective arrangement rendering evaporation from the imer tissue more difficult. (5) Another structural adaptation to drought-was the-increase-of vascular bundles and especially water vessels. The former showed a tendency to coalesce into wooden cylinders near the periphery of the stems. (6) Increased development of the root system in comparison vith the aerial organs indicated a tendency towards higher absorptive capacities, useful in drying soils. (7)_ Volkens found in xerophytes water storing structures such as short tracheids or living cells (idicblasts) in the epidermis of inner tissues. ‘The latter often had a mucilaginous content. Such tissue elements reached large proportions in succulents of salty habitats, but were found also in hypodermal cell layers of W5/039.41NS/Ab/ii5 - page 5 (8) Tnvestigating the epidermis of xerophytes, the author met in dicots with a tendency towards straight, instead of undulate redial valls, a fact corruborated ‘by Lothelier (112). (9) More important was the statement that the mmber of stomata per surface unit vee Often increased, instead of reduced, under ary conditions. This was difficult to interpret in an era in vhich reduction of the stomtal number was invariably regarded as an efficient means for restriction of transpiration losses. ‘This observation by Volkens was underlined by Maximov's too severe criticisn of classic views,. But Volkens understood quite well that the total area of the Pores, as well as their reaction to light and other external influences were rather more decisive for water loss, than their mmber per surface unit considered 80 often as being a too far trusted indication of presumptive transpiration intensity. (20) Volkens was unable to decide whether the resinous substances present, e.g. Delow the epidermis of Tavernia aegyptiaca, or the milky juice in desertic asclepiadeceous shrubs like Leptadenia pyrotechnica, offer these plants any advantage in their struggle i@ is e problem still open to discussion, (11) On the other hand, he found early lignification in xerie shrubs which we might now interpret as an alternative to turgidity of fleshy tissues. (22) Volkens insited already on the"chamical changes of the vacuolar sap" as ‘being an important factor in drought resistance. In the days of Pfeffer" *Qgmotische Untersuchungen", it seemed only natural to appreciate highly the increase in the omotie water absorbing capecities resulting from a rise in concentretion of cell solutes. = (13) More surprising for those early days is the recognition of fundamental differences in the adaptive capacities of various plant groups. One taxonomic group vill be cageble to develop thickened leaves or stems, thus reducing surface with respect to volume, another will develop hairs or strongly ramified roots, while still another group, unable to develop such structural peculiarities, will react to drought in a more primitive manner by shedding part of their structurally unadapted leaves replacing them or not by smaller ones. Such reasoning is fundamental for the understanding of the fact emphasized above that so many different types of xerophytes exist at a one and same place. ‘The oftéi cited treatment of xerophytica by Schimper (195) added little to the list of structural adaptations to drought outlined by Volkens. This femous plant geographer insisted on the principle of reduced transpiration under conditions of water atress and developed the'much-debated idea.of "physiological dryness" of bogs.and salty soils. Kerner von Merilaun (89) offered a different Interpretation of sunken stomata and stomata bearing grooves on leaves d by hairs; He considered ‘the heire as useful. to avoid the blocking of the stomatal catioles by water in rainy periods. Yet, he aia not deny thetr importance for the reduction of yater losses and the protection against overheating of plants groving in lofty alpine altitudes like hair-covered Drabas and Edelwei: um 1); but these plants bear hairs on their upper surface. Kerner ‘lem very W3/039.42. .¥S/AZ/415 - page 6 thoroughly insisting, among other aspects, on the fact that the reduction of leaf surface often means narrowing, resulting in Linear’ shape, a fact corroborated by Cannon (34) for plants in the inner desert of Australia. later authors, such as Neger (143) who, like Volkens, well trained in the eqerinental methods of exact science, added the following items to the list of xerophytic characters: (1) radisation-reflecting, glittering leaf surfaces; (2) water retaining properties of mcilaginous substances, such as pentosans extensively studied and possibly overestimated by Mac Dougall and Spoehr (116) working in the early years of the present century at the Desert Laboratory of Tucson (Arizona); (3) sun-rays evading position of leaves, rising upwards or hanging in a vertical downvard position during hours of intense radiation end heat; (4) production of easily evaporating arometic oils, an adaptation considered as useful by certain modern authors such as Heilbronn (64, in marked contrast to others, such as Audus and Cheatham (11). He also insisted on devices for the absorption of moisture. from the air, a subject which Volkens treated earlier. The latter demonstrated the functional importance of the cells surrounding the bases of epidermal hairs for this purpose. Neger's book is still highly interesting and in many respects up to date; the author combined the qualities of an excellent Plant geographer, physiologist and forester. Another xerophytic character: the dense arrangement of the aerial shoots, vas already mentioned by Ascherson (7) and cited by Valkens (252) We find it in leafless desert shrubs such as Zollikoferis arborescens ani in cushion shrubs, like Anabasis aretioides, both studied by Harder, Filzer and Lorenz in the Sahara, (60) Similar pulviform plants are widespread in sub-tropical and tropical mountainous deserts, for instance Salicornia palvinata in the Andine "Pumas" (Cabrera; 30). We find them also in the Irano-1 ‘Wontane flora (Acantholinon, Astragalus) as well as in the South African Namib Desert (Mesembryanthemim fibuliforme) studied ‘vy Marloth (119). This morphological character is not only useful for the restriction of transpiretion, as found by Filzer (82 ‘but might alternatively be interpreted as an adaptation to potent radiation (Zalensky, 258) end high winds. Both are reduced by the dense arrangement of ramifications which in many cases develop naturally vhile in others they are the consequence of bruising by grazing animals. Eppes of eropytts. Tha aforenintloned cherecters can be coxbined in various manners resulting ‘inite types of xerophytes which have been arranged by ‘various authors according to physiognamical, ecological or phytogecgraphical Principles, Leaving aside short-lived annuals and phreatophytes (ground water Plants), we can distinguish the following main life foms: (1) bulbous or rhizomatous geophytes; (2) evergreen sclerophylls; (3) woody plants shedding leaves or Yeplacing them by smaller and often more xeromorphous ones in the dry season; (4) leatiess, non-succulent brooms like Retama, Genista, Calli ‘and horsetad1- like plants, such as the Australian Casuarinas; (5) plants with succulent leaves, stems or roots;. (6) plants suffering far-reaching dehydration called resurrection Plants. The last mentioned are extreme “troporhytes", i.e. plants changing aspect end activities with changing dryness and wetness (or temperatures) of seasons. Modern an of these life forns have been presented by Huber (75) and by Killian and Lenée (95). : : W8/059.41WS/AZ/¥15 ~ page 7 Ti, Roo? sysTas 5 of ration and Often the degree of resistance of plants to desiccation is a function of ‘the penetration of their roots into shallover or deeper soil layers. If, in early sumer, annuals dry up in the mediterranean basin, flat-rooted grasses, erucifers, composites etc. stccouh first. . On examining plants with leaves in fresh condition, like carrots, dum rigidum, Ononis Hammel, aoe tints ‘that they possess tap-rocts or relatively deep penetiseins aystens. Of course, this rule holds true only if periodic rains vet the sofl to: depths of more than a few inches. The lover and the more scattered the precipitation, the more it is kept in the ‘uppermost soil layers, Accordingly, desert plants are often extrenely shallow-rooted (Cannon; 52). a Weaver's classical studies on the rooting habits of plants of the North American preirie (24%) have shown that to the East of the 100th meridian, i.e. in the true prairie, roots of a large proportion of the plants penetrate deeper ‘than five feet, reaching sometimes thirteen. This permits them to exploit the deeper soil layers after the upper ones:have dried out. The contrary is true of the poor. short grass prairie of vestern Kansas and of Colorado where an intermediary root type prevails and vhere out of eight species examined by Weaver'only one, Peorelea lanceolata, penetrated to more than five feet. In this region, the is t a “widely spreading sbsorbing system in the surface sol". Under full desert conditions trees and shrubs grov mainly in rumnels vhere vain vater collects at least occasionally. If there is any perennial vegetation between the system of these amaller or larger “oueds" or "wadis", their root systas mot be flat and far-spreading, in agreement with the shallowness of rain penetretion, Such flat root systems were found by Cannon in Arizona(32) and in the inner desert. of Australia (34), while Evenari (39) made similar observations in the Judean Desert. : ‘The depth of root penetration proved also decisive when the North American short grass prairie experienced its famous drought ordeal, 4.e. about 193! Albertson and Weaver (3) found that. "bur oak” (Quextus” mac: .). survived vhile most other species dried up. This vas ccomeqisact oF It extensive, deep penetrating root system. Walnut trees, Gleditechia, Maclura also survived, owing to their deep rooting habit. A paradoxical experience was that poplars ried up in desiccated river beds vhere their roots ran nea? the surfece es an adaptation to voll seration.in norm times, but. survived at higher elevations where the roots hed elready ‘to 9 feet in five years. Correspondingly, Mucliar and Weaver (139) atated that pbs deeper rain ani foots bed penstrated the detter seedlings of short prairie grasses survived under confitions of drought, — ‘Thus in their experiments 90-100 per cent of the Bouteloua ilis and Buchloe dsc’ oldes left unmeered for 1h 20 17 days survived: TF thelr Fe roots fad Pere Goll wettened to ten centinctres;” but survival. was 17-50 per cent only vhen the soll was wettened to less then 7.5 centimetres. W5/039.41.HS/AZ/M15 ~ page 6 Tamerous, though possibly not alvays well founded examples of extremely deep root penetration in desert sands, depressions or wadis have been published. Huber (75) mentions the case of Tamarix (or rather Acacia /James; 81 p.1‘ roots found et 30 metres’ depth when the Suez Canal was dug. observati ‘became known from the basin of the Caspian Sea. Vassiliev (231) found roots of Salsola richteri apd arborescens, species of the sandy desert of Kara-kum, penetra ting to opie of several aetress ime majority of the Peammophytes, however, formed extremely long, horizontal roots in the moisture preserving intemediate zone of the dunes which extends from one metre's depth downvarts. In Ammodendron conollyi, the roots reached a length of 20 metres, On the western shore of the Caspian Sea, water relations of semi-desertic plants vere investigated by Kusmin (102) on the Apsheron peninsula. Here, roots of Alhagi camelorum reported also by Keller as very deep-rooting, Medicago coerulea and Glycyrrhiza glabra grow originally vertically domnvards. They branch Gat only at depths bet 10 and 15 metres vhere they strike aquiferous layers. Grape vines bebaved in a simtlar manner. Analogous observations have been made ‘by Stmueli (201) on Alhag! maurorum and Prosopis farcata in the Dead Sea Desert. Their roots penetrate to cc ‘metres. A deep-rooting desert shrub with remarkable biological properties is also the conifer-like Welvitechia mirabilis groving about the 20th parallel in the South West African Wamib Desert. Herre (73) indicates that it strikes underground water in old river beds at. depths of 12 and even 18 metres, The fact is emphatically denied by Walter (241) who found the roots penetrating to 1.50 epproximately, tapping capillary, instead of true gromd vater. Dr. M, Henrici vhon ve asked which author was right, wrote to us that probably both authors were justified each in describing his observed facts; that in spots where veter from heavy mists condenses in flat depressions of a plain (vlakte), she has not seen roots deeper than 1.50m, but in dry water courses or canals, the rooting might ‘be deeper (letter dated November 4 1958). ‘The easy water supply from deep aquifercus layers results in low water deficits of Alhagi camelorun, even during the hot hours of dry sumer days (Kusmin). TE te aloo veapousibie for igh transpiration figures of Acacia tortilis and A.spirocarpa in the southern desert of Israel studied by Zohary and Oreban (264). Tie authors ‘found a root extension of 1.50 in depth and at least 4 to Sm in ‘breadth in the runnels where these trees ‘grow in a region obtaining only about tvo inches of rain. The well belanced water supply even allows those Acacias and certain Tamarix of the. Negev Desert an uninterrupted cambial activity throughout the year (Fehn, 43:hh). Deep rooting of woody peremials in periodically drying- out water courses is also common in the Rajastan Desert (Sarup, 189). Morello indicates it for Lerres divaricata Zuccagnia punctata, Acacia visco, ete., in the ary North-West of Argentina (136;157 I) and Vassiliev for Smirnovia turkestana in ‘the Here~Kum (231)..A favourable vater balance of Aristida pungens and Pennisetum atchotomun sf — is connected with deep root into fe 105). Under less aria conditions deep rooting, rendering water easy and Permanent, has been described by Ravitecher Ferri and Rachid (179) forthe "Campos Cerrgdos", 9 savanna type of Brazil. -The shrubs or small trees in question are “pyrophytes", i.e. plants resistant to fire as e consequence of thick barks, woody underground stems called “xylopodia” or succulent storage roots. They 5/039.NB/AZ/415 - page. 9 easily sprout from buds surviving after fire (Rawitscher and Rachid, 181). This ‘biological group, represented by Jacaranda decurrens (a eae), Craniolaria integrifolia (Pedaliaceae) and Ati exis > @te., contains types with rene! > penetrating roots 0 10 or Li and in the case of Andira Spec. even to 18 or 19 metres. They spend water freely in the ary season Possess neither thick cuticles nor experience high water deficits - a situation which seems to qualify them as pseudo-xerophytes, Besides erosion furrows, valleys and depressions, roots coricentrate in the main water storing layers whose localisation depends upon the soil texture, ~ structure and precipitation. Glendening (52) studying perennial grass on the poor pastures of Arizona, found that the depth of penetration of tiie primary root’ and the zone of origin of the lateral ones clearly reflect the depth of rein penetration. Morello (137,III) found in semi-desert dunes roots concentrated between 10 ana 40 centimetres' depth. Even the few deep rooted species as Atriplex and Larrea divaricata kept 60-70 per cent of their absorbing fouls ie tite lye Variation in the depth of root penetration and ramification depending upon rainfall has been found by Boyko and Abraham (26) in Artemisia herba-albe a characteristic species of oriental steppes, At the eremic boundary of its occurrence, the short primary root branches out alreaty at a depth of 2-5-cm; this results in an extremely shallow root system, With higher precipitation, however, the roots go down to 0 or 50 cm penetrating into vertical fissures of ‘Limestone end remifying only in the rock. ‘The last-mentioned behaviour coincides with the rooting habits of the thorny @varf shrub Potertum spinosum hich covers many squire miles of poor rocky stretches in Palestine's hill country. Litwak (109) recently established that the vertical tap-root of the seedling,-which may grow down to 40 cm length in the first cedeon, later on branches out into the fissures of the underlying rock. Here it uses the vater stored in the soil filling out those cracks. The species grows poorly or is completely lacking on flat sdils covering unfissured rocks. Under mediterranean sub-humia conditions or in light soils, the water- “storing zone is relatively deep’so the roots can penetrate to several metres Yelow the earth's surface as ve established for mediterranean oaks, pistacia (1553156) and pines (147). But under arid desert conditions where flat rooting becomes the rule the plants need an increasingly wide area for their subsistence, and this is the more pronouiiced the shallover the soil and the harder the substrata. In sands only can we therefore expect reldtively deep rooting as @ consequence of a high permeability and low storage capacity of this substrata (Kusmin, 102) (Waiter,239) (Weaver, 24h p.79). In heavy and hard soils of rocky deserts, flat or very flat rooting becomes the rule, as established by the stuiies:carried out _ ‘by Cannon (31), Stocker (215) and Evenari (59) in deserts of three continents and recently corroborated ‘by Morello (137,I) for the poor steppes of Argentina(157,I fig. 22), Here, divariceta usually a phreatophyte, forms horizontal roots six metres long already in the immediate neighbourhood of a drying river bed, but sends its roots don to 30 to 50 cm only. Extremely long horizontal roots reaching as much as 88 feet are developed by Prosopis spici; & psammophyte of | ‘the Great, Indian Desert (Sarup, 189) while the root system Alum dumosuii studied by Evenari (39) xeaches a radius of four metres. This ir W5/059.41S/AZ/435 - poge 10 that the surface covered by the roots of one specimen of Retama roetam may be not Jess than 40 square metres, while Morello (137, II) even indicates 100 to 150 for ‘pushes of larrea cueifolia and divaricata growing on the high plateaux (mesetas) of aria Argentina. Similar conditions. prevail with wild fruit trees-in- Asia. Zalensky (256). studying root penetration of Pistacia vera and almond trees in the mountain steppes of Turkmenistan - with 320 mm of rain end extremely ary sumers - found flat rooting, in marked contrast to the behaviour of the almond tree with higher rainfall in Palestine (Oppenheimer; 146), One of the pistachios possessed strong lateral root-branches running to ten or twelve metres from the place of their origin. 2. Shape of root systems ‘The structure of root systems long stulied by agronomists and foresters, ‘begins to consolidate in our days as a special branch of plant ecology called "rhizology" (Shalyt, 198). In general, we distinguish sympodial or fibrous root systems, videspread in grasses and pains, and moncpodial rot systens with a central, vertical tap root and secondary remifications... The latter type, very widespread in dicots, can be deeply penetrating, with short and sparse ramifica- tions o shortened with stunted main root and strongly developing lateral branches. If both portions of the system are well developed it is Cannon's (32) “general type". However, a superficial system results if the tap-root is at an early stage arrested in its grovth or if_it decays. The general type-ts well adapted to fumetion in both the humid and arid seasons. The greatest pover to exploit the sot] moisture and accordingly drought resistance result when the strong horizontal remifications originated in the uppermost soil horizon send down vertical branches called "suckers", or in Germn Ablfufer. This is the case of Scots pine in Europe and of camildulensis.* An analogous type of grovth has recently ‘deen described jon, ) for Banksia marginate in Australia. A specific tendency towards deep rooting prevails in certais isla exops such as true and sweet clovers (Melilotus) and especially lucerne - for which Rawitscher (176) indicates a depth of 20 metres - ani in treés, like walnuts and pear trees. Plants of arid zones do not differ fundamentally in the types of their root systems from those of other zones; however they seem to be very plastic in their adaptability to aquiferous layers. A typology of root’ systems - founded on his ow observations in the Near Bast - has been developed by Zoheary (261; pp.209-213). In the deserts, tap roots are often deflected fron their vertical course by ‘the dryness or compactness of the lover soil layers. Their deflection favours the formation of lateral roots originating on the convex side of the resulting curvature. ‘This brings sbout “bipartite” root systems ruming horizontally in opposite directions. Such is the behaviour of Suseda tice (Evenari, 39) end of Casuarina pusilla (Specht and Rayson, 211). Alternatively, the tap root ney renin verticnl but otunted or dying back from its tip, ao denonstrated by Morello for Atriplex (137, TIL) while strong or mmerous lateral ramifications developing neat ies orieha in invigorete and form the msin root system. * Unpublished observations in sandy habitats of Israel by Oppenheimer and Kaplan. ws /059-42NS/AZ/415 - page 11 Flat rooting is the rule also in cacti often cited as an example of desert plants rapidly absorbing the moisture.of rare and not deep-penetrating showers. Of course, this is not equally true for all cacti, as was already stressed by Cannon (31). This pioneer in the exploration of rooting habits in the western deserts of North America found that the giant cactus (Carnegies ) posscnced deep-penetrating roots in addition to’ superficial saci He-orgues that ‘this circumstance, which evidently protects the tall plant against being blom over, explains its absence on the flat soils above impemesble "caliche” layers of ‘the Arizona desert where, on the contrary, the flat rooted Echinocactus wislizenit is found. A case similar to that of Camegiea has been described for de Tarilla" in the province of Tucumn (Argentina) by Morello (135); it is Trichocereus teracheckii which has both spreading and descending roots. The ‘latter, useful both for water absorption and anchorage, reach down to 70cm. - The reasons for the develorment of roots running at shallow depths of about 5 to. 10 em below the surface are probably complex, Aeration, nutrients, moisture and temperature relationships, inner correlations as well as specific genetic Properties mst be considered. Boyko (25) interprets the phenamenon in deserts as an adaptation to subterranean dew condensation in a layer 3-8 om deep, after previous studies (Boyko end Abraham; 26) had suggested that Amaranthus blitoides growing in Jerusalem as a ruderal plant, vould dry up in July if there were no nocturnal distillation of water vapour from deeper, moist and war to upper, dry and cool soil layers. This lasting source of moisture, instead of erratic dev, is assumed to explain the subsistence of this species throughout the Jong, rainless sumer. Dev absorption from desert sand by shallow roots is also assumed to occur in Bromus rubens by Killian (94). 3, Pop-root relation It 4s a well established fact that under dry conditions plants develop relatively more roots than shoots and leaves.. Preponderant root growth takes Place in mmerous xerophytes already in the stage of germination. The reasons axe of a hereditary character. Thus pine seedlings studied by Nobbe ¢f. Neger (283; p.514)) under the humid conditions of Germany developed much stronger root system than spruce or fir vhich are less resistant to drought. Later on, physiological conditions play a decisive part; smaller inner water deficits in ‘the groving timof roots than in those of shoots can be expected during ary spells. ‘This will shift the balance of grovth'processes in favour of the roots. On the- other hand, strong light often combined with drought will depress shoot growth and favour accumilstion of ample carbohydrate reserves and possibly root forming substances in the shoot. . Such conditions may be responsible for the surprisingly great disproportion Detyeen roots and shoots existing in several plants of high mountain deserts of Pomir. . Here, the weight of the former surpasses that of the latter 30 to 50 times (Sveshnikova end Zelensky, 221,p.255). ‘The principle of low top-to-root relationships in dry regions vhich can be evaluated as the ratio of leygths freah or dry weights, is clearly pronounced even under moderately Gry conditions. —It was also established Zor the succulent Kalanchoe aegyptiaca by Migahid (127) who found the R/T ratio nearly twice as high W5/059.42.NS/AZ/{N15 - page 12 at 35 per cent soil moisture than at 65,.80 or 95 per cent. But it mst not hold true for attreme desert plants, as etressed by Stocker (215; p.109) who found the root systems of such plants even less developed than those of humid countries. The reason might be that an overdeveloped root system would threaten the existence of the plant by loss of vater to the soil if the roots are not extrenely well. protected against desiccation by impermeable sheaths. Indeed, Magistad and Breazeale (118) demonstrated that rooted cactus joints exposed to an sir stream in oven-dry s0i1 lose. twice as much water from their roots than by transpiration from their shoot. On the other hand, the leaves or assimilating shoots, handicapped ‘by unfavourable conditions of carbon assimilation, almost the whole year round would probably be unable to supply a large root system with food, and consequently Investigations recently carried out by Zobary end Orshan (263) into Zygophyllum dumosum, a succulent, shrub of the Near East deserts, suggest that such Teasoning [. ‘This plant is flat-rooted (0-50 em) during the winter months, ‘but penetrates deeper with short-lived, thread-like extension roots after exhaustion of the vater reserve in the upper soil layer. It seems that water shortage does not permit this plant to maintain a large root system throughout the year and that its extension becomes possible only after accumulation of plastic reserves in the favourable spring season. The changing relation of top to root with declining precipitation is mst Pronounced in the American prairie. Albertson (2) finds the grasses of the short gress prairie only three to five inches high while the roots penctrate considerably deeper. In one case Bouteloua with shoots 1/2 foot high had roots 5 feet deep. Corresponding figures vere 2 1/4 and 9 feet for the deeper rooted Psoralea tenuiflora, 3 and 12 feet for Kuhnia glutinoca, 1 and 6 feet for Solidago glaberrim, ‘T/R length relations were 1:3.5.to 1:6 in the xerophytic grasses as against 1:3 in the more mesophytic "big bluestem" vegetational type. Mueller aml Weaver (239; p. 397) conclude: "The vigorous roots of shortgrass seedlings early develop extensive absorbing surfaces and are well suited to supply large quantities of moisture to the tops”. A critical study by Schopmeyer (194) of the properties determining the drought resistance of shortleaf pine, Pinus echinata, in comparison with loblobby pine, Ps taeda led to the conclusion that the greater development of the root system with Tespest to the shoot 1s the-decisive factor... The usefulness of-such a éisproportion is often quite cbvious, as in the case of Lotus cretious studied by Killian (92) on and dunes. He found a close correlation between survival of seedlings and the length of their tap roots, However, the stronger development of roots must not of necessity mean a large extension in depth or breadth. Instead, we often find a, greater density by increased secondary and tertiary remifications. Hereby, a better utilization of soil water is possible if it becomes immobilized in the wilting range of soil moisture, As is well known, the vater melon, colocynth and other cucurbitaceous enmials ove their drought resistance mainly to their intensive rooting babit. Already Miller (130) was able to explain the higher drought resistance of sorghum WS/039-42., 19/aZ/425 - page 15 in comparison with maize mainly by this principle. The former vas found to possess approximately twice as many secondary roots per length unit of primry root as hed the.mize plants, Thus their mass development was mich greater ‘though they 41d not. extend beyond the limits of the maize roots. ‘A special feature of root systems of steppe and desert plants is’ the form- tion of very long and delicate "rain roots" developing within a very short tine, sometimes a few hours, after a rain shower. Stocker who observed then in Egypt (215; p. 113) as well as in the Hungarian alkali steppes (217) regards them as highly characteristic of these habitats, Their function doubtless. is the temporary increase of absorbing surfaces for a far-reaching exploitation of thé rapidly appearing and again dwindling water resources. As soon as the soil aries, ‘the rain roots disappear and the permanent skeleton roots are tightened again by dorky sheaths the strong development of which is no less characteristic of desert plants (Volkens' 233; p.26). : h, Velocity of xoot growth ‘The ready onset of root growth in arid plant formations has been observed by several botanists but little information seems to exist about the velocity of root growth in dry regions. Oppenheimer (147) indicates 5 to 6 mm per day for the primary seedling root of the Aleppo pine and 3 mm for the oak Quercus cailipeinos: In a.root tunnel arranged at Jerusalem (148) the tap root of Stone pine Fspiaee) seedlings grew as mich as 8-10 mm per day, secondary roots of Pinus halepensis 6-9 mn. . These figures are small in comparison with those reported by Cannon (33) of a Progopis velutina seedling which grew 51 mm in 12 hours in a root thermostat et NOES "However, in spite of their relatively lov grovth rates, the tap roots of the oak seedlings studied by Oppenheimer (148) in an organ-pipe like arrangement consisting of narrow, vertically arranged, wooden boxes of increasing length, reached 55 to ebout 0 cm during the first year of their life end 1.60 m in the second. In tropical sevannas, the strongly heated soil during the dry season has caused the origin of woody plants with horizontal or vertical subterreneah stems and/or roots. Part of these plants bear the character of subterranean trees with only relatively short, delicate leaf-bearing shoots projecting above. the ground. Such is Andira humilis in Brezil studied by Rawitscher and Rachid (180). Smaller perennials have relatively large, woody underground stems: such as the afore- mentioned | iziapotis at or near the surface, Jacaranda decurrens and Creniolaria Antegrifo: 4, 175). Again others like Cochieo: ‘insigne have fleshy tap ote ot attaining sue metre'a depth (Ravitecher ant Reskid, Ter ye Solna hf studied by Birand (19) in the neighbourhood of Ankara has a 8: ‘tap root. (This author also describes the root systens of various perennials escaping death by deep penetrating branched or unbranched. root systems). Succulent plants living underground with their leaves hardly touching soil evel axe Lit salicola and Nenanthug vittatus studied ty Walter (240). The roots. of. these pl 3S emerge at 5 ‘em Below'sotl level. The root-stock of certain South American @varf palms belonging to this group of underground plants, such as Acanthogocos specs, can descend, after germination, to 50 cm depth.(80). ‘That of Attalea exigua, the leaves of which project but little above the ground, goes dovn even “ems. . Specht and Rayson (211) describe similar adaptations w5/039.41.NS/AZ/415 - page 1 from the sclerophyllous heath of South Australia. Here, the caudex of Xanthorrhoca australis is found between 23 and 45 cm below the surface and the creeping root- Stock oF 1 ichotoms at 15 cm. ‘These adaptations, as also the succulent Lignotubers Sa ‘by Beadle (17), are very important in connexion with fires sweeping many tropical and subtropical areas. The water and food storing sub- terranean organs with their renovation buds are situated deep enough to protect these plants against intense heating caused by sun or fire. Thus these phyrophytes escape destruction vhile other species without such buds perennating underground are alnost invariably annihilated (Specht and Rayson, 120)... In Brazil, these | “fire plants” of the cerrados of Sao Paulo seem to have immigrated from the drier interior of ‘the conbinent after destruction of the primeval forest which set in already in pre-Columbian times. The presence of subterranean vater-storing organs in many epecies of the South African veld has been stressed by Henrici (66; p.627) who speaks of a "tremendous development of water-storing root system, as subterranean stems and roots." Among these, Walter (2h1) investigated Pachypodiun succulentum and Kleinia redicans, plants with underground tubers, for their water @. Yor South America an excellent description of plants with succulent underground stems or roots has been provided by Cabrera (30). This author. deseribed nev species from.the dry Puna, to the East of the Cordillere:’ Perezia burckartii with a turnip-like water storing root and very low T/R ratio, Trichocline mcrorrhiza, stemless with a thick, spindle-shaped root and Forephyllue grenoohllim, a dver? shrub vith a thick and woody root (29)... These are ti the knowledge of the Andine hizb mountain deserts called by A.V. Humboldt’ (76) “a: solitudines, tempectatun vicissitudinibus mire Obnoxiae". The life forms is Fegion have tee ‘efficiently sketched by Neger TiS; p.196-202), Water-storing fleshy roots were also sean by Vassiliev (231) in the Kere-kum Desert. Burkill (28) discusses the armoured tubers of Dioscorcass 5. Root penetration into rocks A new field of research concerning root life in semi-arid zones lies in the study of roots' extension and ramification into rocks, Already Kusmin (102) tablished that porous stones reseubling aquiferous rocks in texture absorbed vater in the wet season, releasing it again to invading roots in the ary sumer, The present author observed active penetration of vertically growing pine roots Anto soft limestone rock if the covering soil layer had dried out in the early sumer. Zohary and Orshansky (262) found that certein limestone rocks had a surprisingly high water storage capacity and that dvarf shrubs of the mediterranean “rocky heaths" or"phrygana” (Pritzel, 124); such as Varthemia iphicnoides, Stachys Beleestine ena Potonomma syrinon ovuld grov on the vere, unt arisenes ‘rock, their root: ving as sy grew in soil. Oppenheimer (151) explains the surpris- ingly high transpiration rates of Quercus ithaburensis in localities where no or Little soil cover remains, with root grovtli in the unfissured bed-rock; this conclusion was recently corroborated by Halfon - Meiri (59). He was also able to demonstrate the root growth of pistucias (P-palaestina and Lentiscus) and Quercus ealliprinos in solid rock of either porous or. softened character (155, 156). . The ‘Toot follows here in the footsteps of physical vesthering. It disintergrates the calcareous and possibly other rocks, but it is evident that this capacity is a privilege of highly adapted pioneer species excreting relatively concentrated acid compounds. The importance of these species, including probably buckthorns, wild fig and Capparis osa, for the resettlement of sites badly eroded by higher Plants, Sear ceasidorabies This te ‘quite obvious in the case of the lentisk WS/039.41.NS/AZ/415 - page 15 which, after its firm establishuent in one place, develops long, straggling extension branches in all directious.. These strike roots at nodes so that one shrub which originally germinated in a tiny depression of a rock, may finally cover an area of dozens or even a hundred square metres. Another case of a mediterranean dvarf shrub penetrating into soft rocks with its mmerous fine rootlets 1s Thyms cepitetus groving in Israel as a rock plant. on extremely shallow soils shove soft Line ant pontatonee, the plant has no tep- root: the rock’ storing some water even if covered only by a very shallow soil layer, makes deep penetration of this dwarf shrub superfluous (Litwak, 109), -The same author stated that he had seen rocts of cistis also growing in such soft, unfissured rocks. 6. The influence of soil texture and structure and of chemical composition As in nore humid regions, the soil texture and structure play an important part in root development in semi-arid and arid countries. Mrs. Halformeiri (59) working with the present author finds that the Palestinian Valonea Osk Quercus iteburensie develops much denser root systems rich in absorbing rootlets if it ‘grows in heavy rendzina or basaltic soil, than in sandy, less fertile soil types. ‘These very Obvious differences can in our opinion hardly be attributed to the higher water storage capacity alone, but rather to grovth stimlating substances Imown to occur in the soll organic matter and to mycotrophic mtrition. On the Other hand thie’ presence of concentrated soluble salts precludes root growth, and numerqus. reports, among which the recent one by Karschon (86) insist on the fact that in highly saline and dry deserts leaching out of salts from runnels and wadis is a primary condition for the establishnent of higher plants. 7. Anatomical properties of the roots cf xerophytes When roots enter the secondary stage of develorment, the drying up primary , . cortex usually exfoliates and disappears. In desert and steppe plants, hovever, it often persists. The peripheric layers may becane hard and corky protecting the plants against the pressure of bard soils ( lon articuletum; Killian, 93) and desiccation. The imer cortex instead may become spongy (by perforation of the disintegrating cell-walls) forming dead sleeves. This is thé case with 5 numerous South African grasses, Gerbera, Commelina etc. studied by Henrici (67) who describes its function as that of a vater-absorbing and storing tissue, an interpretation undérlined by the experiments of Killian (94) and Lemés (105). While the former found that such roots of Bromus rubens were absorbing imediately upon contact with liquid water (12 per cent of their weight by capillary suction), the latter found that the dead roots of rtium vere strongly hygrostopie absorbing 100 per cent of their original wei moist air. They conduct Lguid water eastJy, like mosses, both at their periphery and in the central conductive bundle: Walter (241) on the other hand, insists on the isolating properties of these sleeves protecting the living central tissues from the desicoating influence of the surrounding soil. Roots ‘of sandy habitats in dry regions possess often very dense and long root- hairs agglutinating sand by more -or less mucilaginous secretions of pectic substances, Killian (95) considers such permanent roct-hairs appearing near the apex of roots of Haloxyion articulatun as functional even in very dry soil efter prolonged drought. ws /039.RS/AZ/415 - page 16 we GERMINATION AND ESTABIISEMENT OF SEEDLINGS 1, Difficulties in naturel reproduction As is well mown, natural propagation of plants by seeds is strongly handi~ capped under arid conditions because seed ripening on the mother plant might be incomplete the germination percentage low, owing to hard seed coats, low soil moisture, hard non-cbsorbing soil crusts (Beadle, 17), high concentration of soluble salts, etc. But even when a seedling has struck a root, it. will ary up rather than get fimly ebtablished, oving to Viciesitudes caused by irregular rainfall, destruction by animals, mechanical lesion, wind erosion (Shreve, 202), covering vith mid, competition of older established plants or of quick groving annuals choking them out (Beadle, 17). Also, the intense heat and radiation in the arid zones 1s much more dangerous to tender seedlings than to better protected mature individuals. Foresters know that natural roproduction of forest trees proves often ineffec- tive in semi-arid regions. Reports such as those of the Forest Administration of Cyprus (229) offer mumerous illustrations for this argument. Even artificial establishment of forests by reseeding is frequently ineffective and succeeds only sfter planting of sturdy, vell lignified nursery material grom in containers. Often tree seedlings, for instance oaks, are very sensitive to dry air, in contrast to the mature, deep rooted plant. The fact that the majority of. exotic woody plants introduced into Palestine rarely reproduce from seeds without human help, was stressed by Oppenheimer (149). ‘The drier the region, the more deéert like the vegetation, the worse are the chances for survival. The classical investigations of F. Snreve in the Arizona Desert (203) showed that the Giant cactus (Carnegies sigartes), though able to germinate on dry sand without a drop of water (( iy > Teaches an extremely low percentage of reproduction. Shreve calculates that on a northern slope with total of 240 specimens, there were only three to four survivals per year in the 45 to 60 years old plants, one among the 50 to 40 years old and none during the last ten years. (The abnormal distribution of ages was apparently a consequence of a more than usually dry period or of increased soil erosion). Similar.conditions prevailed with Parkinsonia aculeata (20h). For this tree, the "palo verde", only 0 to 3 per cent of the seedlings survived after three years while later, after this critical period, the death rate becomes negligible, | It.is true that part of the skeleton branches will desiccate in dry sumers, but the rest regenerates, a behaviour also reported by Arcichovsky (5) for the saxaul desert forests of Transcaspia. During four years, Shreve aii not find a single young plant a Encelis farinosa, a sumflover-like perennial widespread in the Arizona Desert 7 Miss Henrici (69) reports on the extremely poor regeneration of Karroo bushes from seeds. From 100 germinating seedlings hardly 5 per cent will reach maturity. In a closed veld reserve, she did not, see even one young bush during the first six years! w8/039.41NS/AZ/415 - page 17 Also Morello (135) insists on the lack of seedlings of perennial plants in the poor bush’ steppes of the region of Tucwman, due to the fact that the various favourable, factors, required for survival through the first critical years, rarely concide. A most unfavourable factor in the district. is the late summer @rought occurring after riperiing of the seeds of the creosote bushes, Larrea divaricata, Monttie aphylia, and others, While in the South African veld, Favourable tanperatires for germination in Spring axe often ineffective, since ‘there is no rainfall, better conditions prevailing in autum are frequently Zollowea very soon by frost which kills many seeds even before their ripening Gienriet, 72). : In an earlier paper (66; I) this author, describing conditions on the’ Amoedvlakte, a poor steppe to the North-Zast of the Karroo Desert, had already described the complete absence of young plants, especially perennial grasses, "the most striking feature of the yeld” where seedlings mist be both drought and frost resistant. Thé very low natural reproduction is a striking feature also on the lofty” heights of the cold Pamir Desert, according to Sveshnikova (220). Low precipita- tion often does not moisten the soil sufficiently for seed germination. and, even if this happens the hard climatic conditions are responsible for subsequent enormous losses of seedlings. Under less extreme conditions, the high mortality of seedlings is @ phenomenon occurring in more or less frequent dry yedrs and periods only. Irregular and inadequate rains following germination caused in the Dark Island Heath of South Australia, the death of many seedlings in'winter.’ From seedlings other than Banksia ornata, 60 per cent were killed by summer drought in 1950, the first of their life, but only 12 per cent in 1951 (Specht, Rayson, and Jackman, 212), Investigating conditions in the North American prairie, Blake (20) found a nearly 100 per cent mortelity in grass seedlings germinating in spring, due to heat, drought and the competition of well established, older peremniais. The rocts of the latter form a dense mat through which the seedlings penetrate only with great difficulty. , Chances for survival are better in winter when seedlings, having germinated in autum, are protected from the cold by dead leaves and snow. Modern research has tried to analyse the causes of this tremendous waste of energy in ineffective seed production: On the other hand, it bas endeavoured to elucidate the adaptive chatacters enabling the seedlings to overcome the perilous stage of their early life through useful adaptation, and also how this could be furthered by sulteble treatments. We sball discuss first the eforementioned problen: the causes of the large lot + Blake (20). found the germinsbility. of seeds: defective with prairie grasses in dry sumers only. ‘Henrici (71) insists on the bed quality of seeds produced by wilted @mases puch as.Themeda, while Syeshnikova (220) found no seeds at all in Gry years, i.e, in the Pamir Desert not more than 30-40 um of either snov or rain. If seeds developed at all, gexminability vas rether good. A high percentage of empty onde ims found, only 4n fou dyen{és,, 1s Keantholtam Sepeibitses Even . leguminous seeds (Astragalus: is) gexmina’ wi ‘special treatment. Sane. of the: species "very low temperatures approaching zero. © Thus ~ the unfavourable results are essentially due to external factors. ws/039.42WS/AZ/415 - page 18 ‘Tn mmerous cases, hard seed coats impermeable to water have been found responsible for low germination capacity. This was the case with Cercidiun floridum and Delea spinosa studied by Went (249) in the southern desert of Califomia. “Here, germination can take place only after mechanical lesion of the seed coat r in certain cases the epicarp) by sand or rubble transported by water or vindborne, Of course legumes contribute a considerable contingent ‘to these cases, as in xerophytes of the Kara-kum Desert (Sveshmikova, 22), Trigonella arabica (Koller, 96) and Colutes istria (Koller.and Negbi, 98) in the tegar Bosert ‘of Israel. But here, also several species of Convolvulus evinced the same. property (Negoi, 18). Difficulties can be overcame in this case by mechanical lesion or pretreatment with concentrated acids. It seems that in certain cases seeds of other propagules might rot even in deserts after drenching rains. Koller (96). assumes that in certein psammophytes Like Calligonum comosum, the fruit of vhich is protected by a dense cover of prickles, tkts aivangament prevente exaggerated motstening of the frutt which ‘vould rot without ample oxygen. This is still hypothetical but fits in with Kerner's (89) ideas regarding the water repelling function of hairs on the lower stomata-bearing face of leaves, 2. Germination inhibitors A quite modern trend of research was initiated by Went (249) who found - in agreement with indications fron Henrici (72) from South Africa and views expressed by Cavazza in Italy (36) - that often no germination will occur in the Californian Desert before a considerable rainfall, 10,60 millimetres or more, bas drenched the soil. This was interpreted as due to inhibitors present in the Dropagative bodies: seeds, fruits or aggregates of such, They delay germination or hamper root develoyment of seedlings if they are not removed by abundant rains. Went's hypothesis, since corroborated by experimental investigations, disproved the present author's early assumption (1h; p.308) that germination inhibitors in fruits were to be expected in rainy climates rather than in steppes and deserts where the lack of water alone seemed sufficient to prevent premature germination. Went's investigations offer convincing proof thet many desert seeds possess germination inhibiting substances, and subsequent research in Israel underlines his results. Pectis sa (Went; 249) a composite, requires at least 10 ma of rein for germination ‘Temains dwarfed if the seed is leached insufficiently. Many seeds in the Death Valley behaved similarly and even died from growth inhibition if rains had been too light. In phytostat experiments, Went (250) found earlier germination and stronger early develoment of desert plants, such aa Filago, Boutelous, Gilia if previously washed by heavy artificial rain (300mm). This, however, proved noxious in other cases (Tillsea, Eriophyllum, Plant ete.) Generally speaking, no germination of winter annuals takes place in the Death Valley if rainfall is less than 60 m.(Went and Westergard, 253). Soriano (209) working with Went as Pagadena, found 10 to 15 mm of rein optimal for germination of desert plants, such as ‘lum ambi gum and species of Pee! vhile others, such as Ameranthus sas dest results with 355 mm. Activated coal improved germination suggesting that adsorption of inhibi- ‘tors can replace leaching to a certain degree, Poorer results obtained after W5/039.41NS/A2/415 = page 19 heavier rather then optinel rain might in our opinion, possible be interpreted a8 the result of leaching of minerals which are necessary for initiating enzymatic activity, from the éeed or of the germination stimulating substances, from the soil. A bint that germination inhibiting substances may be present also in the Black Heath of South Australia can be found in the statement by Specht, Rayson and Jackman (212) that for satisfactory germination, at least one inch of rein must be stored in the sot. ‘In:Ierael's deserts, germination inhibitors called also “blastikolins" have deen found in the fruit valves of fm dumosum, the bracts of Atriplex adie and of A.dimorphostes ant fe the perteoniad leaves of Runex roseus (kolier, 96) while im Colutea istria studied by Koller and Negbi (98) a substance delaying the growth of the seedling 1s present in the outer hard seed coat. Soriano (209) found no influence of rain on the easily sprouting seeds of ‘the "Rose of Jericho" (Anastatica), Cutandia hitica and the colocynth. On ‘the other'hand, 100 to 200 um of seis iuproved Fecilts considerably with seeds of the cruciferous Erucaria boveana and Carrichters annua. ‘The removal of inhibitors (mstard oils?) becsme apparent in Erncaria also by longer stems and better devel- oped leaves of the seedlings. In non-desertic plants of Palestine, Zohary (259) had earlier established that intact fruits inhibit the germination of isolated seeds of Onobrychis evistagalli. This is-in‘accordance with investigations by Cavana CE) on the hear related, cultivated seinfoin ~- also @ me@iterrancan species. The author found thet germination of the seeds is checked not only by the mechanical action of the legune hindering imbibition, and by. lack of oxygen, but also by inhibiting substences delaying the emergence of the radicle and hampering its development. Fruit valves of Sinapis alba contsin a blastokolin effective on its seeds (Zohary, 259; Sroelov, Vahl (230) detected germiation inhibiting effects in the fruit of Poterium spinosum. Blastokolins in glumes and pales of Aegilops, etc, which are parative only at high temperatures, help to prevent premature sumer germination (Weisel and Adler, 238). Tt seems therefore thet. germination inhibiting substances are widespread in arid end semi-arid hebitats.. This is further underlined by Went's observations on the death of seedlings in the neighbourhood of desert and chaparral bushes in California. arrea seedlings die in the neighbourhood of adult Larrea shrubs (253) & case: where -- by mearis of substances excreted by its roots - a plant sees to Kill its own offspring appearing so near to its own self as to threaten its’ fiture existence. Ina later article (252), Went reports on the bad influence of Salvia Mellifera on the germinating seedlings of Adenostm. In this case the alien species hada stronger adverse effect than Adenostoma bushes. Went's surprising observations can be classified under the concept of “allelopathy ~ the’ influence of one plant on the other? developed ty Molisch (13l).if we are not dealing rether with substances diffusing from-dead leaves and other components of litter shed by the chaparal shrubs. Such litter inhibits germination of both Salvia mellifers and Rhus leurina (Went, 252). - * : W5/059.42NS/AZ/415 - page 20 Similar observations have recently been made in Israel by Donner (38). Continuing our om studies (153), he analysed the reasons of the striking differ- ences in the composition of vegetation under trees of Quercus ithaburensis, as compared with their open surroundings. He found that fresh or the last season's oak leaves not only obstructed ‘the development of seedlings mechanically but contained also germination inhibiting substances. Germination of grasses (Hordeum Spontanerm, and bulbosum, Avena sterilis, Dactylis glamerata), legmes jun clypestum, roussaeanum) and others was strongly delayed by extracte from these leaves. The latter exerted also @ grovth-depressing influence on young seedlings of the mentioned species. As is often the case, the author found sometimes a stimiating, instead.of an. - inhibiting influence. This may be explained either by dilution of the inhibitors or possibly be caused by grovth promoters produced by fungl or other micro- organisns. We wish to add here in passing that Shreve (20%) found the influence of the Shade of mother trees negligible with Parkinsonia seedlings. On the other hand, Sarup (189) is responsible for the statement that the shelter fron the blazing sun offered to the young seedlings in the Rajasten Desert has @ beneficial effect, as would be expected by most botanists and foresters. Humphries (178) found the Disnaga (Echinocactus wislizeni) growing to maturity only if protected by shude of mesquite Bushes etc., in its seedling stage. The chemical analysis of blastokolins end related substances has recently made progress. Jn a paper sent for publication Lerner, Mayer and Evenari (106) show that in lum dumosum and Trigonella arabica osmotic effects of soluble salts: (NaCl) are complement ether soluble organic inhibitors. . In the latter species, the ether extract of fruits contained coumarin an unsaturated lactone kmown for its inhibiting effects on germination but apparently not found earlier in fruits.as a blastikolin. 3. . Speed of germination snd growth of the radicle Among the adaptive characters developed by desert plants, speed of germination after effective rains seems to be of utmost importance if the seedling is to survive in the subsequent dry period. Quick emergence of the redicle is followed in Fopguicre oplenaers studied by Livingston (110) ty its penetration into deeper layers which do not ary out easily. Tn the fifty years and more elapsed since Livingston's classi¢al studies ‘these observations have been corroborated by many others. Sveshnikova (220) found that up to 75 per cent of seeds of Eurotia ceratotdes germinated within 18 hours vhile 45-80 per cent of Artemisia skorniakovi sprouted within two deys: Also, Went (249) established in his experiments that desert seeds germinate easily. The same is true of theearly rot develomment of the seedlings. Quick penetration of the primary root into the soil before considerdble development of ‘the shoot was described ty Blake (20) and Plumer (171) for mmerous plants of the prairie, by Glendening (52) for the "tanglehead" grass (He contertus) of Arizona,’ ond by went (S49) for perennials and shrue of the Saute Calivorai ‘Calirornian Desert, like Dalea spinosa. This plant developed roots 40 em deep before the WS/039.41S/AZ/415 - page 2 shoot reached a length of more than 3 ci, In this case deep penetration is fevoured by water storage of two reiny seasons per year in the semi-desert soil. Herre (73) grew Welwitschia seedlings in three feet long drainage pipes to suit their deep rooting Habit, While Wells and Shunks (246) indicate that seedlings of Quercus catesbaci had roots 5 1/2 inches deep in their two leaves’ stage. An evea greater disproportion was found by the present writer in seedlings of Quercus cal os (147): while the young shoot was 2. cm long, the tap root attained 35 mm. On the other hand we are informed of the failure of species mable to get seedlings established because ‘of the lack of quick growing primary roots. This is the ease for prairie grasses such as Agropyron smithi1 and species of Festuca, Elymus, etc, mentioned by Plumer (171). rello (135) insists on the fact that ‘roots of shrub seedlings often lose contact with the retreating moist layers of the soil during the autumnal dry season while Waisel (236) explains the failure of Tamarix ephylla seedlings in sandy deserts by insufficient root grovth; in his experiments roots bad not reached more than 12 cm in length in 80 days when ‘the ssind dried out. . 4, Pretreatments In experiments’ aiming at developing agricultural production in groundwater free desert areas, as those recently undertaken in Israel, speeding up of germina- tion by pretreatments plays an important role, Such experiments sumarized by Negbi (142). include emong others: soaking for removal of inhibitors, vernalisation at cool temperatures, illumination of seeds inhibited by darkness, ¢.g. Oryzopsis miliacee and thermoperiodic stimilation, An interesting example is that of Penicum antidotale an Indian perennial tried out for pasture under conditions where not only quick desiccation of the soil but also the formtion of a hard crust makes sowing a gamble. Here pre-scaking at 6°C. and subsequent drying offers good prospects by accelerating the germination process. Afterwards the seedlings have a feir chance to got a firm stand in the field-before a new crust cuts them off from the underlying wet layers, as it often occurs also in the bare "scalas” of the inner deserts of Australia. Glendening (52) finds establistment of grasses in Arizona negligible without sone covering of the seeds by Miter or soil in analogy to conditions im. deserts with impervious soil surfaces. 5. Temperature effects ‘It has long been known that seeds are, as a rule, adapted to higher or lower ranges of temperature, in agreement with the occurrence of the species in.cool or hot regiots. Under subtropical conditions this results in e well distinguished yeanly cycle of germination of sumer annuals sprouting in spring end winter annuals sprouting in eutwm. Such changing aspect: of vegetation is very pro- nounced in countries like Israel in the weed flora of irrigated fields and gardens. It is even more obvious in the semi-tropical deserts of California with their two- seaéonal type of rainfall, Went (249) found species with tropical affinities, Like Aristida ‘adacensionis, Boerheavia, Amaranthus and ia germinating only after sumer others, like gilts, > ‘Lepidium and ‘belonging to’ the stock of the northem’ ‘zone appear efter ¥ reins only. . Perennials and shruba germinste, as 6 rule, in summer only. . Went considers this pronounced thermal adaptation as typical of desert plants. -Infeed he considers that eurythermic species germinating both in sumer end vinter are WS /059.41W3/Az/¥15 - page 22 not typical desert species. Surprisingly enough, this neutral group contained thermophilous plants such as Datura and Cucurbita sta. ‘The present author feels that Went's conclusion might prove premature, investigations might show that the above temperature ranges have little to do with dryness, but essentially with adaptation to the main thermal belts of the globe, as mentioned by Went himself. Actually we find similar adaptations in subtropical forest trees. Oppenheimer (147) found the germination of the-Aleppo pine taking place normelly detween 4.5 and 20°C. but rapidly declining at higher temperatures (25-26°), ‘This behaviour has recently been corroborated by Magini and Mugnaini (117) and found true elso for Pinus pines (optimm 17.5°C.) and - vith little modifications (optimm 17.5°C.) - Sy Wagaaint 140) for Pinus pinaster. All these mediterranean pineo ere thas adapted to germinstion in the Soot seaicH and Will not sprout at 26° or more prevalent in the mediterranean basin in the hot sumer months. ‘This probably reflects the thermal adaptation of section Pinaster of Pinus the species of which are concentrated in temperature and even cold zones. On the other hand, eucalypts, if som in Israel in winter, will not germinate at all or at best very slowly being adapted to a wamer environment. In spite of possible differences in the interpretation of the facts, it mst he edmitted that the thermal adaptation of the eycalypts suits the conditions of a semi~tropical comtry with prevalent summer rein, It should be added that Gondelmann (55) found various mediterranean geophytes groving wild in Palestine, i.e. Urginea, Scilla, Pancratiun and Muscari, also germinating under cool winter conditions only. The sane holds for Trigonella arabica (Negbi, 142) and probably for mmerous other geophytes and winter. ennuals. In contrast to these mediterranean and oriental-steppic plant types, seeds of desert plants in Israel seen to be a little more thernophilous. Koller's (96) data suggest that they germinate satisfactorily in the interval between 15 and 30°C. Same prefering slightly cooler, others like Citrullus co’ germinating as Koller says with great difficulty as long as the ¢ ook coat is intact - higher temperatures. Different day and night tenperatures stimilate germination of Panicum turgidum (Koller, 96) while Went (250) found thet asa rule, some desertic summer California developed best if the nights weré cool vhile winter anmals prefer cool day and night temperatures. While -karroo bush seeds germinate in many cases at 20° as vell as at 30°C., many will germinate ‘better and earlier at the lover mit of this interval, some even preferring 15° (Henrici, 69). Indeed this author recently recamends this lower temperature for best practical results with "good karroo bushes". (72). A very striking phenomenon is the stimlating influence of intense heat on many seeds of bushes or trees in semi-arid comtries. It is partially responsible for widespread germination after fires, e.g. in the Australian vattles. In their dry state the seeds are less sensitive to heat than would be expected. - Went finds (251) that seedlings of Ceanothus leucodermis and others appear in Sarge mubere in thé. Californian chaparral, after s fires ; after a fire. The seed of the latter is not killed by 120°C., but will produce a meximm number of seedlings afterwards. A corresponding optimm for dry heat pretreatment for Adenostoma fasciculatua and ¥8/039.42.15 /az/5 ~ page 25 Styrax oficinalis is not less than 150°; seeds of the latter species still peau 50 per cent seedlings even after heating to 140°, Specht, Rayson and Jackman (212) found that species of Helichrysum and Stipe macalpinei germinate only after fires. The high figures of germination often established after fires are, however, frequently a consequence of an increased release of seeds from hygroscopic cones, as in Aleppo pines in Israel and other conifers. The follicles of Banksia ornata releases seeds only after a fire which also favours release of seeds from woody fruits of Casuerina pusilla and Leptospermm (Specht, Rayson and Jacloan, 212) Went (252) further insists on the fact that lessened cotpetition and the destruction of germination inhibiting litter favour germination after a fire while the Australian authors insist on the importance of mineralisation of nutrients useful for the development of the seedlings. 6. Protective adaptations of seedlings Tn numerous cases, germination is immediately followed by the formation of water-storing tissues that guarantee the seedling's existence in the subsequent ary period. The seedling of ere _splendens stulied by Livingston (110) undergoes early thickening of Soo SE is further protected by cork defore the epicotyl emerges, This preparatory stage of development lasts two to three weeks. Conversely the seedlings of ia versicolor develop turaip- like storage roots lacking in the mature Sonastian (oamseas SI) Cannon, 31), The tiny Othouna minima and related species resist drought by aid of Aisproportionate ‘subterranean tubers (Compton, 37). The primary root of Que: catesbact, growing on sand in the coastal plain of North Carolina, siete Beteuing "elougate- ‘tuberous at the upper end functioning as a water storage organ" (Wells and Shunks, 247), Similar to it is the root of Tit jis ipecacuanhsae forming storage tissue throughout (for more data on a eee re see paragraph TIT, "Root systems"). In other cases the seedling's root possesses contractile properties diewing the young stem downwards into the soll. Such a behaviour, earlier described by | Thoday (223) seems to be typical of all South African species of Oxlis with endosperm containing. seeds (Salter, 188). By the geophilous movement, axillary and‘other bulbils sitting on the stems are transferred to the depth required for their further development. . Anbrosinia bassii, a mediterranean Aracea studied by Killian (91) forms both tubers ani contiactile roots in the seedling stage. Th hie study on the kinetics of geophytes, Galil (47) devotes a chapter to. the ‘movements of mond and dicot seedlings into the soil during the first stage Gf their development, “The vital part of the seedling bearing the growing paints af root ani shoot. ts passively pushed dommard by & ‘promoter! Be latter my st of 4 tion of 1, as ‘Allium, aris consist of “basic ‘ports only cotyledon, aie a and Sternber; or of the tube-shaped united cotyledohary’ . ne te pe ee P [elds at of the mediterraneén region in Palestine, the promoter can’ reach a length of 30 cm or more. Consequently the corm developing foun the hypocotyl below the prouoter, originates fron the beginning ‘ws/039-NS/AZ/415 - page 24 at this depth and 4s protected from eradication by Arab and ordinary European ploughs. Nothing except the blades of the two cotyledons appears above the soil in the first season. : In other cases the downward pushing growth of the cotyledon is continued by a stolon-like caudex, as in sharonensis, or "guiding" roote came into action. These fmction as pacé-mikers. Eventually they are obliterated inside, forming hollow tubes. . Through these the firet leaf, shest carrying the spical meristem of the seedling, grows downwards. This type of early-development, is found in species of Colchicum, Tris and others, Finally the downvard growth of the cotyledon moving the spical meristem of the shoot forward can be replaced in a latter stage of develorment by the pulling movement of contractile roots, as in the above case of Oxlis and in species of Bellevalia, Gladiolus segetim, Leopoldia growing on the mediterranean territory in Palestins. . Problems of seed dispersal ‘The biology of germination of plants groving in the Near East covers a vast field of phenomena connected with the dispersal of fruits and seeds. The scope of the present article is too restricted to allow a detailed presentation of: the facts and ve can mention here only two basic contributions: those of Murbeck (11) and Zohary (259). These authcrs have shown that, in contrast to earlier assumptions, very often no dissemination of the propegules to distant places takes place in steppes and deserts. On the contrary, the young plant germinates in the immediate neigibour- hood.of the mother plant, and arrangements such as heavy or sickle-shaped fruit, sticky seeds or even active burying of the fruit in the soil by the mother plant (geocarpy, amphicarpy) ensure that the often rather restricted suitable “living space" is not lost. Part of the propagules at least sre not equipped with accessories facilitating their transport by vind. This-is the case with certain heterocarpic composites producing achenes with scarious ‘cups or stiff bristles, jastena of feathery pappus. A nev, amphicarpic bean has been described by Burkert 27). 8. Differences in germination readiness and capacity Recent research has demonstrated that morphological differences in the shape of the propagules (fruits, portions or aggregates of such, fruits plus bracts, seeds, ‘etc.) are very often matched by pronounced physiological differences. ‘The seeds germinate either immediately upon their ripening or after a longer or shorter - delay. . Zohary (259) found very pronounced differences in speed and energy of germination between the basal and apical portions of cruciferous fruits of the Brassicese and Cakilinae tribes. In white mustard, seeds included in the apical portion of the fruit did not germinate at all. SroBlov (214) investigating the uetter further, concluded that germination inhibitors are present in Sinapis, Erucaria, Matthiola preventing early germination of part of the seeds. recent works by Koller (96) and Kedman-Zahavy (83) have contributed additional fects. Atriplex dinorphosteris, investigated by the former, bears fruits surrounded by et ‘or bumped bracts. Seeds of the latter germinate after sae delay, 5/039.42§8/82/415 - page 25 possibly not after the first, but after subsequent rains, In Atriplex rosea studied by Kaduan-Zahavy, we find smooth and wrinl:led propagules the former containing brown, the latter black seeds. Black seeds germinate considerably later than brown ones: their majority will not germinate at all in the first 7 season. Similar results have been obtained by Beadle (18) with Australian species of the same genus. Such differences are interpreted by certain authors as “germination regulating mechanisms" fulfilling thé function to “scatter germination in time” or at least to restrict 1t to a definite combination of external conditions. Such mechaniems would help to prevent the disappearance of annual species. This would indeed be the consequence if total germination of all present seeds were produced by a favourable coincidence of conditions followed later by conditions unfavourable fer further develonment. While we do not deny that such interpretations might be Justified in certaia cases, we feel nevertheless that this attitude tovards action and interaction of multiple, often highly complicated, external and internal factérs Opens the door to futile teleological speculation in others.’ In the present phase of knowledge it seems preferable to collect facts and to refrain from premature interpretations regarding the advantages of only this or that combination of e.g. bard, impermeable seed, light or darkness, inhibitors, etc., for the survival of the species. Such interpretations ‘are apt to lead astray especially if they are made on the basis of laboratory experiments insufficiently checked by field observation. We therefore restrict ourselves to the mere statement that frequently seeds of dry regions germinate only partially in the first season after their maturation. Zohary (2593 p.58-59) found that of the dispersal units from numerous legumes such as Medicago, Astre: Onobrychis, ete., only one or tvo seeds germinate in the first winter, the rest in later years, The same holds true for the "synaptospermic’ dispersal units of Aegilops and the eremic Pteranthus echinatus. Several authors find that seeds germinate only at certain seasons. ‘Thus Blake (20) met with a pronounced peak’ of readiness’ to sprout in spring end a seisnéry ons in eutum in plante of the prairie. Tn Atriplex diaorphostegia, Koller (96) found seasonal differences ‘in germinability, ‘are possibly the consequence of an endogenous rhythm. sis miliacea from Galilee gerninates only in part during the first winter the rest 70! the next. (Negbi, 12). "Tt mst be admitted that in these cases a spread in tine exists. It can de the consequence of external or intern&l factors; often temperature might’ be _ decisive, in cohination with seed properties. Renrici (69) found few of the tested karroo seeds germinating promptly. The majority either requires long periods for after-ripening or germinstes well only at certain seasons. Thus seeds of muiltifidus will begin to. germinate only five months after maturation reaching 6 peak of readiness to sprout after 18 to 2k months... ‘Tetragsnia arbuscula germinates only after an after-ripening period of two years. Welafrida genicalata will reach its peak ~ probably: due ‘to a combination of after- periodicity - after 18 months, The great influence of the “latter factor can be i}ustrated by Tetra, arbuscula. At its seasonal peak, it produced 86 per cent germinations:’ three days while ‘three months ‘vere required to obtain 5 per cent geminition vhen sown at an unsuitable deason. W5/059.4.NS/A2/415 ~ page 26 Also some desett annuals from Arizona studied by. Barton (15) do not sprout without after-ripening. dium lasioc: reached its optimm of readiness after one year, Streptanthus ieus "26 months. 9. Mass production of seeds On the other hand, many annuals seem to counterbalance the danger of extermination in dry habitats by profuse seed production. Went (250) stresdes ‘the fact that sometimes after rains in the Californian desert many thousands of. seedlings are found on each square metre, Such density of ephemeral vegetation 4s one of the many paradoxical phenomena in deserts. In contrast to the ‘behaviour in plant coenoses of a more permanent character, these plants do not arrive at a developmental stage where serious competition leads to the elimination of the weakest - which are thus prevented from reproduction. Went confirms - earlier statements asserting that even the tiniest ephemerals arrive at setting fruit and ripening seeds, just as Erophila verna on roofs and walls in humid regions mentioned by Timirlazev (225) and others. In spite of thése tentencies tovards mass production of seeds and germination under often very definite conditions only, prospects of’ effective propagation’ in arid regions are not bright, especially for perennial vegetation. Accordingly. we find in deserts and semi~deserts a pronounced tendency tovards vegetative propagation by sprouting from roots, rhizomes, tubers.and bulbs. Henrici (66) insists oa the ready shooting of apparently dried up grasses taking place “with Incredible speed” from the margins of old tufts in the Armoedsvlakte, and on the fact that during the season of sumer rains "the bare spaces between the grass tufts became populated with mmerous creepers espec‘ally Convolvulacese and Cucurbitacese", ‘Albertson (2) found that-after the great drought buffalo grass Yepopulated the free es in the prairie by its creepers. Atriplex halims, studied by Evenari (39) in the Judean desert, sends shoots ‘rom ite enceping root-stock to the surface, and Cannon (34) insists on the frequent occurrence of stolon-like: organs in the desert of South Australia. The surprising density of geophytic vegetation spreading underground and sprouting everywhere from the bare soil after the first rains is a comon experience of people living in mediterranean and adjoining steppic comtries. ‘These examples which could be multiplied ad libitum, illustrate the great advantage of established vegetation over new-appearing tender plants in arid surroundings. 10, Light ‘The influence of ight on germination of arid’zone plants is complex varying with wave length and temperature.. Therefore we refrain from including these “photoblastic" reactions in the present report, mentioning only the interesting observations by Henrict (69) who found karroo seeds in’ general adapted to germina- tion at low light intensities or even in, the dark Ike those of Petragonia arborescens, The fruits of this species "bore themselves into ‘soon, fter wettening apparently thus avoiding thé depressing influence of light on mamisatton, Specine of Pegolevtie ani Peronia Hhbw « atellar bebsrioar. ‘he author recamends covering ted seed in-the veld with branches to improve results. . W5/059.u1 :ws/az/45 ~ page 27 Yor further light effects the interested reader is referred to Evenari's recent review (41) and to the work of his associates: Koller (96), Kadman-Zahavi (83), Negbi (142) end others. . V. LEAF DROP AND SURFACE REDUCTION If a plant does not obtain enough water to balance its umavoidable losses, it often will reduce its surface by shedding leaves. As a rule, the process Degins from the oldest leaves and progresses tovards the apical meristems sparing ‘the youngest leaves. As Pringsheim (173) was able to show in Pfeffer's Institute, water is transferred from the fading leaves upwards, especially in succulents. Mothes (138) added the observation that concomitantly proteins are broken up and the resulting products (asparagin, glutamin) also translocated to the shoot apex. Even potassium, magnesium, phosphoric acid and other easily mobilized nutrients are vithdrem and translocated. Often we find development of short latéral shoots (brachyblasts) with tiny leaves from the axils of the dropped old leaves. The new small and thickich leaves can be kept alive and turgid even with a much reduced water supply from the roots. Reduction in muiber and size of leaves is a highly effective means of edapta- tion to occasional or periodic drought. Its importance in phylogenetic develop- ment during the tertiary epoch is stressed by Axelrod (12). Fading and shedding of old leaves during unusual dry spells is a widespread phenomenon during dry summers even in the temperate zone. In the mediterranean region many deciduous trees afd shrubs, as plims, almonds, apricots, hawthorn, buckthorn, Melia azadimchta etc., show premature defoliation much more than evergreens which part ith their leaves rather reluctantly since they serve also as storage organs. They are drought-enéuring rather than drought eveding. The mediterranean shrub Anagyris foetida and Erodium gt im ‘in the desert (Evenari, 40) are leafless in sumer bearing leaves and flowers Say ia the rainy season. ‘The rocky heaths of the mediterranean shores and the Near East abound in low shrubs with superficial root systems which shed their larger leaves of the rainy season replacing them regularly by sualler sumer leaves, As examples we cite ‘Thymus capitatus, Varthemia, Salvia trilobe and Poterium spinosum, ‘The same phenomenon is found in the desert where leaves are often replaced by brachyblests, reduced to stipules or shed completely. Examples ere Heliotropiva rotundifoliun, Suseda asphaltica, Atriplex halimis (Evenari and Richter, 12]. We further mention species of Teliaathams, ‘Tdnaria aegyptiaca, Astragalus sus and others, Nitraria retusa, Zilia ea, Lycium arab: ‘and Ochradenus baccatus - the two latter studied Ty aay Oat Orta (26h) - and inmmereble others. ‘While there can be no question sbout the effectiveness of surface reduction es a means of adaptation of transpiration to the dwindling water supplying power of the soil, little quantitative data regarding the phenamenon are available. . Jerdent (82) studying Ononis Jeiosperm and Orshansky (identicel with Orsham; 158) Ononis natrix shoved that in these species the sumer leaves are gradually reduced in size, pessing in the latter case through @ series trifoliste-wfoliate-petiolate vile stipules of very reduced size are nreserved even in late summer. Jardeni found the vater content of the sumer leaves, as related to dry weight, reduced ‘by more than two thirds, This enhances further the effect. produced by reduction W3/039.41us/az/ia5 ~ page 28 in number and size. ‘The extent of the assimilating organs (stipules plus petiole, as compared with the same plus leaflets) is in Cnonis natrix reduced by ebout 85 per cent according to estimate from Orchansky's plate. Shmueli draws winter and sumer leaves of Alhagi maurorun from the Dead Sea shore. Here the surface of the sumer leaves finsily is no more than 2 per cent of the winter leaves which @issppear in July while the thorns increase in inverse proportion. In Nitraria retusa, large spring leaves dominating till Jue are replaced by axillary clusters ‘of two to seven thickish sumer leaves. These are gradually reduced in size the smallest and innermost only remaining til1 the winter rains. Zypogte/lium duogwn etudied by Zobary an Orsban (263) also sheds its leaflets: nothing ‘petiole remains. The transpiring surface is thus reduced by 87 per cent, while evapotranspiration per 100 square metres of the sparse etn dumosi association drops from 6,377 kilograms in winter, including March, 5 In the hot and dry season (April to September) i.e. by more than 95 per cent. A survey of reduction in weight of the last season's spring flush has been carried out by Orshan and Sand (160) in shrubs of the three main climatic zones of Israel. Teble 1 sumarizes the esscntial results calculated by the present author from their unpublished data, Mediterranean small shrubs reduced their green weight by one to two thirds in most cases, mainly by changing their leaves. The same is true of the psemmophyte Artemisia moaos} in the Negev Desert where {t enjoy the relatively favourable moisture Suditions of its sandy habitat. Other semi-desert and desert shrubs pay a m:ch heavicr toll reducing the weight of their young shoots by more than one half in all cases, but sometimes by tvo thirds, three quarters and more. Table 1 - Shoot weight of currant year's growth cycle at the end of the dry sumer, in percentage of spring weight. (calculated fram data by Orshen and Sand) I. Mediterranean: Judcan hills. 1957 1958 Phlamis viscosa 5. 52 Gistus vitlosus a 59 ‘Teucriua polium 4s 43 Scrophurlaria xanthoglossa 50 4h ‘Thymus capitatus 36 50 II, Ireno-Turanian, Semt-desertie: Revivim Artemisia herba-alba 28 26 Noce mucronata 2 ° III. Seharo-Sindtan, Desertic: South of Bersheva Haloxylon articulatum 28 % Zygopaylima dumosum wy meee Spec. 4s 3B .. to., Grier locality Ea B Artemisia monosperma (psemmophytic) 45-52 . & 5/059NS/AZ/N15 - page 29 Tt should be added that to the summer of 1957 folloved @ normil winter to that of 1958 a relatively dry winter where rains after February were negligible. While this had a decided effect on absolute shoot. weight, the table shows that it produced no marked effect on relative losses in weight which seem to be relatively independent from the preceding precipitation in e one and same climatic region, if ‘the rainless period is very long. In an earlier paper (159) Orshan had stated that the dry weight of leaves per unit length of shoots is reduced during the dry season, as follows: Poterium spinosum by 84.5 per cent, Artemisia monosperma 76 per cent, Helianthemun Hlipeicun 62.8 per cent.’ “Having foun Be difference in the tranapisation rates of sumer and winter leaves calculated per unity of fresh weight, in the period of transition, he feels convinced that with certain desert plants of the Near Best, reduction of leaf number end mass plays the decisive réle in water econony and adaptation to the rainless season, while regulation of water loss by the leaves often plays a minor part in the desert, in contrast to mediterranean species. investigated by him. Morello finds both determinants of total transpiration important. Studying the water balance of larrea and other bushes of the “monte” (137; II), he finds reductions in leaf surface between 18 and 80 per cent in the dry season; the higher figures were obtained with Larrea cuneifolia. The dry season lasts here for six or seven months. The coubinad effect of lesf drop and mainly stomatal regulation of water loss from the remaining leaves reduced the total transpiration 8 rather low frections of what had been established in the wet season (see table 2). Table 2 - Approximate calculation of transpiration in the southern district of the "Monte" (After Morello /137,117.) Critical season: 6 months; Rainy season: 6 months. In litres of vater for each plant. Dry season Rainy season Minima. Maxim = Minima ~Maxim = Amual total larrea divaricata 90 300° 1350-1450 1450-1750 uccagnia punctate ho 150 1550-1450 1400-1600 larrea_cuneifolia 90 300 360 150 450-1450, A far reaching reduction of transpiring surface in sumer is found in Pennisetum dichotamm growing in the Sahara. It sheds all its leaf blades. The rydroeconamic effect 1s further increased by the fact that the remaining haulms with their surrounding leaf sheaths transpire per mit of weight five times less than the blades. (Lemée, 105). ws/039-41NS/AZ/415 - page 30 ‘While the data accumlated recently suffice to permit a first insight into the quantitative aspects of periodic leaf drop in arid regions, Zohary (260) hes recently proposed a double system of classification of desert plants founded on ‘the manner and the time of reduction of their vegotative organs. The time- classification comprises only one evergreen category: ‘the Acacia-type that keeps some leaves even in the driest season. ‘The drought escaping Retama-type and the Ellngo-type shod leaves very early, ‘before the end of the reiny season while the yunsea and Reaumuria types keep them till June, ‘The second classification distinguishes types according to the character and Place of the shed organs. Here ve find leaf, shoot, branch and cortex "shedders" ‘among more or less leafy or aphyllous plants, The new classifications were proposed by the author because he felt that Raunkiser's system of life forms is inapplicable to the xerophytes of the Near East. Zohary's system might become more workable and generally recognized if nearly-related types are collected in more comprehensive categories vith short and pregnant Greek or Latin names. Morello (137; II) has added a few more types to Zohary's morpho-ecological classification. A list of Central Asia desert plants shedding leaves or shoots at the Deginning or in the middle of sumer is contained in an article by Sveshnikove and Zalensky (221). Species of Calligomm, Ephedra, Kochia, Artemisia, Selsola and Chondrilla are mentioned (3.258). the erticle sumarizes recent work on xerophytism in the Soviet Union citing, among other works, a survey of “historical- ecological” xerophytic types in the Pamir-Alai region by Grigoryev (57). economy of brooms While the broom-like vegetation type has often been interpreted as an adaptation to periodic soil drought, relatively little physiological research has ‘Deen carried out to substantiate this hypothesis. Investigated in the temperate zone, these plants are not distinguished fron other plant types by low transpira~ tion rates. Indeed, they contain poly- and oligohydric (Oppenheimer, 154) types, i.e. spending either mich or little water; they may be 1so- or poikilobydrie (stable or labile in Stocker's /2167 terminology) in their water expense, as recently demonstrated by Leyerer (108). Their curface transpiration is often vather high, and even if compazed on a fresh weight basis with mesophytes they must not transpire less (Lemée 103), Even under desert conditions a broom like Retama roctam can, according to Evenari and Richter (42; p.348, etc.) evince high ‘ranspiretion rates; one single shrub attaining a water consumption of 1500 to 3500 kilograms per anmm, thanks to its deep and wide spreading roots. On the other hand, Leyerer has demonstrated that Sarothamus sco) reduces its total transpiration by more than one half ty shedding its leaves. In the Algerian desert he found brooms spending already in spring decidedly less water per grame of fresh weizht than leafy plants of the same region (the experinents took place after a rather ary vinter, with little water reserves in the soil). Sarcthnmnus scoparius and Ulex eus - in contrast to Calluma a and Gantsta plicga are able £0 s0ebrict their transpiration effectively ‘their xeromorphous structure (Lemée, 103). Zobary (261) finds even Retama roetam decidedly not polyhydric, and his diagram (p.22h) shows that its transpiration drops sometimes to about 100 mg per gramme fresh veight in an hour. W5/039.h1: m5/A2/IO5 ~ page 32 ‘This is in accordance with Evenari's and Richter's findings who indicate corresponding low figures for. the end of the dry season. Sumarizing present knowledge, we may argue that the brom-like plants of arid end semi-arid zones are able to reduce their water expenditure effectively ‘by permanent or seasonal surface reduction in times of water stress. For Iayerer the hydroeconamic advantage of the leaflescness of this plant group mainly lies in the fact that broans must not spend water as other plants in order to avoid overheating of their leaves in dry periods. Thus the hydric advantage would be indirect, the radiation-biological advantage direct end of primary character. In contrast to such an interpretation and mder the impres- sion of the results of Konis (100) who found the cooling influence of transpiration on leaves in moving air rather unimportant, ve prefer to stress the direct influence of leaflessness for an undisturbed vater economy. We further should Like to insist upon the fact that already Kerner von Merileun (89) had interpreted ‘the broom type of the mediterranean shores as an adaptation to high winds in merked contrast to similar-looking hygrophytes growing in all regions of the Globe /89, vol.I; p.3957. This seems still worthy of eamest attention. Tt is certainly not by mere coincidence that the leafless condition becomes more and more widespread in non-swamy habitats if one proceeds from humid to arid coniitions, It finally becomes dominant, as in the Grest Indian Decert of which Sarup (189) writes: "Most of the plants of the area have no leaves and are mere tvigs". Such are for instance Capparis a and Euphorbia royaleana. In the Kalahari end Karroo deserts 36 plant types are also a Very striking feature and certainly they dominate in the southem sector of the Negev in Israel, one of the driest regions of the globe. Here we find Calli govum cousun, Haiowylon persiom, Eeselicomnicu, Retana recta, Ausbasis articulates, = a wus baccatus wich, even Tumels as they do, are either ‘completely leafless or bear little and small leaves only in the rainy season, Effects of @ reduction by rolling of leaves ‘The effect of rolling on the transpiration of gress leaves has been investi- gated by Lemée (104,105), He shows that the effect is inconsiderable with hygroyhytic types, bearing ample stomata on the lover flank of their blades and possessing monbranes easily perneable to weter vapour. On the other hand rolling of leaves was found highly effective in grasses with a pronounced xeromorphous structure, i.e. thickened epidermis valls reinforced by a ring of hypodernous fibres and few stomata sunken in protected grooves. Such are e.g. Stipa tenacissima (Tschirch, 228) from the Old and St.ichu from the New World (CasteLianos, 35). such grasses growing in the Mediterraneis and the Auvergne, complete rolling reduced transpiration by 46 to 6 per cent. The effect was even more pronounced with Stipe, and Festuca duriuscula growing in the Sshare in complete agreement ‘earlier conclusions by Techirch. Here the reduction produced by rolling reaches 69 to 83 per cent i.e. figures very similar to the above mentioned produced by far reaching leaf drop in other desert plants. If Bennet-Clark cited by Grieve (56) was unable to confirm the usefulness of Jeaf rolling for water econamy, this might possibly be a consequence of the choice W3/059.41NS/AZ/415 - page 32 of plants of @ not too pronounced xeromorphous structure. We camot say this definitely since we did not see his paper. On the other hand observations by Lemée and by Henrici (66; p.605) demonstrate clearly that in many cases leaf rolling of grasses takes place at high deficits only, and the leaf dies before ‘the mechanism can prove helpful. We can confirm this for Srmodon. dactylon. In rolied other cases grass leaves are from the beginning awl-shaped ‘as those of spartium, Aristida pungens and St: "lora (Lemée, 105), even under conditions of a good water supply. ris it would appear unjustified to speak in all these cases of "xeroplastic" (Thoday) adaptation evoked by drought, without critical experimental studies, lenée sumarizes his report by the statenent that effective rolling of grass leaves at sufficiently low water deficits is relatively rere and by no means more frequent in deserts than in less xeric habitats. Vi. GUTICUIAR CONDUCTANCE ‘The water retaining properties of plants living in dry habitats depend upon different mechanisms, such as the vater retaining capacities of protoplasm produced by water retaining substances (mco-polysaccharids or protinaceous compounds), the impregnation of the cellulose framework of the epidermis walls With fat-like substances such as cutin, or the presence of suberized cork layers. ‘The outer epidermis wall is, as a rule, further reinforced: by a covering of pure cutin, as in the leaves of Clivia nobilis (Anderson, 4), and in Agave americana forming the cuticle proper. There are further often additional coverings of waxes known for their pronounced hydrophobic properties or varnish-Like resins widespread in xeric localities mainly of the southern hemisphere (Volkens, 234). The available information about the chemical character and submicroscople arrangenent of the cuticle and the adjacent cutinized layers has recently been summarized by Treiber (226) and Skoss (203). It seems sufficient to mention here that in cutinized cell walls cutin forns a high?y polymerous three-dimensional fremework consisting of cutinic and cutic acid, cutinin, ete., more or less esterified by monovalent alcohols. The meshes of this framework - which often is associated with pectin and cellulose - are more or less stuffed with wax excreted by very delicate plasmatic threads running through a system of very thin pores. If the excretion. is strong, wax will eccumlate es a bloom composed by rods, plates or massive layers on the surface. The excretion of cutin and wax is stronger in sun than in shade leaves. - Water stress increases excretion of cutin, With rising tempersture, relatively less cutin and more wax is produced (Skoss, 208, experiments with Nicotiana glauca). This will exert a strong influence upon permeability to water vapour since the cutin framework alone does not represent a hermetically tight structure. Pectin, also present in the epidermis valls and cellulose, is hydrophilous and considersbly more perneable to water than both cutin and vax. If stomatal restriction of water emense is practically complete, as is the case with certain succtlents, selerophylls and desert bushes in periods of severe drought, the reduction of “cuticular conductance” or, in other words, the tightness of the outer epidermis walls, becomes of decisive importance for resistance to desiccation. It has been argued that no cuticle is absolutely impermeable to water vapour. In mesophytes the cuticular part of transpiration can reach a considerable fraction of the total water loss. Thus Kemp (85) working on the Rhine, found it to be one half of total transpiration in Acer syriacum, a figure closely agreeing with results Ws 059.41Ns /az/415 - page 33 of Mittmeyer (132) who investigated Rhododendron ponticum. In a tropical : sclerophyll, as Coffea arabica, cuLisulay Wranspinetion saounted to ane fourth of the tote In Oleander and laurel, however, which represents sclerophylls of a neditervaycan character, 1t amounted only to 1/13 and 1/45 respectively. Rawitscher end Ferri (178) found in Cedrela fissilis 1/60 to 1/70, ‘These examples illustrate the wide variation of cuticular conductance which plays also a considerable part, in the absorption of dev and solutions by leaves (Arvidsson, 6; Waisel, 255; Skoss, 206). Tlirtel who investigated cuticular transpiration recently (61, 62, 65) found it much influenced by actual acidity and the lyotropic influences of. salts. Cutin seems to behave as an ampholyte reaching minimm figures of imbibition and consequently maximm figures of permeability at a neutral or weakly acid reaction (pH 5-7). In contrast to suberin characteristic of cork layers it is tightened ‘by icns favouring imbibition, the swelling of its micelles exerting a narrowing effect oa its submicroscopic pores, While salt influence might be considered as not uninportant for cuticular transpiration of halophytes - high sulphate possibly increasing it - the investigations of HArtel and Vukovits (63) on the influence of ultraviolet radiation seem also of ecological interest. These authors find a reduction in the swelling capacity of the irrediated epidermis and consequently increased cuticular transpiration. For xeromorphic leaves with strongly developed cuticular layers, dehydration by the air, called “incipient drying” in its initial stage, is of fundamental importance. Glumann and Jaag (49) who studied the influence of changing hunidity on cuticular transpiration in the dark of sclerophylis, found with Rhododendron ponticum end Pimus silvestris irreversible shrinking of the external cuticular — layers. It exertéd an enormous depressing influence on cuticular transpiration. Thus in the case of Scots pine the dehydrating influence of the atmosphere - which reduces trenspiration not gradually but by jumps, and is opposite to that of dehydrating solutions studied by HArtel - reduced original cuticular transpiration ‘by no less than 99.5 per cent. We may assume that the tightening of the cutinized epidermis valls is a combined effect of dehydration and chemical constitution. As indicated above, cuticular wax plates are intimately interwoven with the highly polymerous meshes of cutin macromolecules (Treiber, 226). If the whole structure shrinks in a dry atmosphere, it becomes practically impermeable to water. Ecological. literature offers ample proof of the high efficiency of cuticular restriction in plant transpiration. Weaver and Clements (245) cited by Shields (199) are reported to have stated that cuticular trenspiretion of xerophytes under water stress is practically nil. Oppenheimer (146) found cuticular transpiration very low and soutimes nil in Laurus nobilis, Arbutus andrachne and the Aleppo pine investigated at the ‘end of the dry season in Palestine. He could later (150) ada Syprosens, semperrizens vo the list. | Also the eventos oak Quereus_ cal iprina and even luous Pistacia stina (150, 154) were found capable 6: complete suspension of Epenaplretion tater Severe conditions of atmospheric and soil drought. Very low figures of transpiration meaning the loss of about 1 er cent of fresh vetght per hour vere also found in other mediterranean sclerophylls like olive and carob (Oppenheimer, 146). Leaves of both may sometimes lower transpiration to nil (Poljekoff,'172; Oppenheimer, 150 for the olive.) These W8/039.41,WS/AZ/h15 - page 3% results demonstrate an extreme drought enduring adaptability of mediterranean s trees avd shrubs to arid conditions. Similar cases have been reported by jenari and Richter (42) for Retama roetam and Haplophyllum tuberculatim in the Judean Desert. =o The effectiveness of the cuticular restriction of transpiration is also demonstrated by the results of Pisek and Berger (163) with isolated, heavily eutinized sclerophyll leaves, like Rhododendron and ivy, Norvay spruce and Scots pine studied in the laboratory under moderately dry conditions. Cuticular transpiration vas in all these cases below 1 per cent of fresh weight per hour. In conifers it even approached 1/00-per hour, These results seem to agree with ‘the thickness of cuticles messured by Gdumann end Jeag (49) who found cutinization in Abies sibirica and Pinus silvestris heavier than in Rhododendron, The high efficiency of the cuticle in pines is confizmed by the results of Pearson (166) for species from the semi-arid states of western North America, He estates: “Western yellow pine, Douglas fir, Bristlecone pine and Engelmann spruce possess @ surprisingly high degree of resistance to transpiration when the water supply is reduced to a dangerous level." ‘The latter conclusions. are confirmed by Parker (162) who could keep needles of vestern yellow pine living for tvelve days and of Douglas fir for six days at 40 per cent relative humidity and 25.5°C. Their very low cuticular losses vere covered by the transfusion and assimileting tissue undergoing considerable shrinkage. Less extreme restriction of transpiration with closed stomata wes found by Walter (2h1) in the sclerophylis of the South African "broken vela". The cuti- cular or, to say the least, mainly cuticular losses related to fresh weight amounted with Rhus erosa to 1.5, with Osyris compressa to 1.8 per cent per hour while with Roycna microhpylla and-Rhus burchettii they reached even 10 per cent and more. Transpiration of Cedrela fissilis Jacking pronounced xeromorphous structure can nevertheless be very effectively curtailed when the stouata close. Leaves of that plant studied by Rawitscher and Ferri in Brazil (178) will lose through cuticular transpiration about 0,5-1.2 per cent of their fresh weight per hour. - ‘The authors calculate the daily cuticular loss of a whole tree in the dry season and find 2.5 litres per day when leaves are young, and about 5 litres when they are mture. ‘This consumption can be covered even by the scarce precipitation of the dry months in the region of Sao Paulo. In a recent study Ferri (45) found in the Campo Cerrado of Sao Paulo a maximum cuticular restriction of total transpiration to 1/50 in Annona cariacea while in the semi-arid Caatinga formation in North East Brazil only Spondias tuberosa reached comparable figures: 1/20 to 1/50. In all the other plants studied, the cuticular losses reached 1/10 to 1/3 of transpiration with open stoma. The results suggest that cuticular conductance approaching the zero Jevel is very rare in'the tropics, even in regions experiencing periodic drought. Probably it presupposes either fer reaching and irreversible desiccation of the cuticle or additional layers of vax or resin. the factors exerting their influence upon the efficiency of cuticular restriction of transpiration the thickness of the cutinized leyers and age of the leaf warrant detailed discussion. Kamp (85) established the important fact that WS/039.h1.NS/A2/415 - page 35 in structurally and taxonomically similar leaves cuticular transpiration is more or less inversely proportional to the thickness of cutinized layers (Olea e' versus O.lancea;” Fhamnus alaternus versus Kh. 1aulosi If, however the leaf with the thicker cuticle has a relatively restricted surface and is relatively nore succulent - as is the rule in xerophytes and also in sun-leaves of the same species in comparisoh with shade leaves - its trenspirative capacity is hereby increased and no simple inverse relaticuship between thickness and cuticular vepour loss is found. In these cases there might be better agreenent with the assumption of inverse proportionality if cuticular transpiration is related to weight instead of to surface. No wonder therefore that many physiologists 1tke Seybold (197) and ecologists find no clear cut connexion between cuticular thickness and cuticular transpira- ‘tion (Svenert and Richter, 12). The relationship is further complicated by changes with age. Rawitscher end Ferri (178), Kamp (85) and Mittmeyer (132) agree that older leaves exhibit higher cuticular transpiration than younger ones. Kemp explains the increasing permeability to water with age by a process of solution and disintegration brougut about by atmosrheric factors while Mendel (225; p.77) considers cracking and fissuring as a consequence of repeated moisture deficiency. Such demege is partially repaired by further thickening occurring in ageing leaves of certain species. This will not alvays compensate for the ageing effect if cuticular conductance is measured at high humidities but prove very effective at low ones. While the effectiveness of heavy cutinization of sclerophyll leaves as 8 means of restricting transpiration in times of emergency is underlined by Seybold (ign, Oppenheimer (152) and Grieve (56), others like Ferri (15) and even Morello (137) seem sceptical about it. The latter goes even so far as to” conclude that xeromorphiem has nothing to do with xerophytisu since numerous xeromorphous plants grow in by no means dry surroundings (157; p.356). While @isagreement about the interpretation of this undeniable fact will probably continue Zor long concerning sclerophyls, we did not hear so far of anybody questioning the use of heavy cutinization for preservation of moisture in cacti though even these highly xeromorphous plants have delegated representatives as Eptphylium and Fhipsalis into tropical rein forests. Living here as epiphytes, They enjoy the protection of their thick skin in the rainless periods. A classical investigation into the effectiveness of epi- and hypo- dermal ‘tissues of cacti on their transpiration in the open and in the laboratory has been carried out by Mac Dougal, Long and Brown (114), Experimenting with Echinocactus wisligenti they established that the uprooted plant kept in diffuse light, ‘Lost jaot more than 0.05 per cent of its weight per day. Remaining alive without supply of water for more than six years(!), its daily loss in weight finally dropped even to 0.015 per cent per day. The cuticular transpiration of this cactus calculated per unit of fresh weight was therefore about a thousand times smaller than that of heavily cutinized sclerophyllous leaves. Among the latter, those of laurel and olive lose about 0.6 per cent per hour, even after closing of their stamata, according to their om unpublished observations. Plat tia, studied by the above mentioned authors lost - if left without water Bualy — atout 0.15 per cent of its weight per day and can keep on without water for many months. As Martin (121) has shown, peeled Opuntia Joints lost W3/039.41ws/az/415 - page 36 in 48 hours no Jess than 87 per cent of their weight as against 4 per cent (apparently with slightly open stomata) of controls. ‘The result agreeing remark- ably well with findings by Trippi (227) exemplifies the overvhelming influence ‘of cuticular or, more correctly, pachydermel resistance, as compared with the rather weak water retaining capacity of the strongly hydrated inner tissues. The latter apparently ewters into the picture, if at all, only after the stored unbound water bas easily evaporated and the micilaginous cell contents of the storage tissues help to keep back the rest. For the structural peculfarities of the cacti and especi- ally their mich developed sclerenchymatic and collenchynatic hypodermal layers, and of other succulents the reader is referred to Boosfeid (22) and to Mac Dougal, ‘Long and Brovn (114). It should be also mentioned that Migahid (127;p.25) found stomatal closure restricting transpiration more effectively in succulents than in non-sueculents a fact demonstrating the high efficiency of their cuticular protection. South African succulents studied by Walter (21) lost in most cases more in weight than North American cacti, Daily losses of. the uprooted plants were ae follows: 3-10 per cent in Kleinia redicans, 1-2.5 per cent in Crassula platyphylla and in Meseub: em ‘and about 1 per cent in isolated EFluteccaices of Alse species Lithops selina lost 0.25 per cent only per day ‘thus approaching the figure for Platyopntia established by Mac Dougal, Long and Broa. Resinous coverings While there can be no doubt that the presence of vaxy Incrustations inend waxy bloom on cuticles strengthen their water retaining capacity, as was recently shown quite conclusively by Skoss (208), the interpretation of resinous, varnish- ike coverings in the same sense has so far found no unanimous approval. Volkens (254) who met with this anatomical character with Leguminosae (Baccharis, Celmisia), Saxifragaceae (Escallonia), Acacardiaceae (Rhus), Compositae (Vernonin) and ‘liacese (Larrea), assimied that these substances - which he found soluble ee ett |, potadslim hydroxide and benzene - fulfilled a transpiration restricting function. Actby (9) studying structure and transpiration of Larrea tridentata, arrived, hovever, at the conclusion that the creosote bush possesses no efficient transpiration restricting mechaniams.. In his opinion this plant evades death in ary periods by protoplasmatic adaptation to high water deficits and vy its ability to recover from prolonged wilting. Runyon (186) confirming the epparent lack of pronounced xeromorphous characters in this plant, nevertheless would admit that the varnish layer should restrict transpiration. This opinion had been advanced earlier by Spalding (210). The careful anatomical (137,1) and physiological (137,II) investigations by Morello seem to settle the problem definitely in favour of Volkens and Spalding, es far as the genus Larrea is concerned of which three species were examined: wcuneifolia, divaricata and nitida. The leaves of the first mentioned species are able to reduce their water expense by stomatal closure to one hundredth of the initial figure in the course of one hour. Larrea divaricata lost under conditions of serious water stress no more in welght than ©> mg per square decimetre im an hour. Such restrictive capacity is also very pronounced with Zuecat Punctata another shrub with resinous epidermal covering. Morello tes that ws/039.41.NS8/Az/415 - page 37 a leaf of this species vould spend all its water in less than one hour vhen its yta are open while in @ closed condition this process would go on for 7 days and Q hours, We thus get the impression that these South American xerophytes with their - as Morello finis in contrast to Runyon and Ashby - pronounced xeromorphous properties, do not owe their drought resistance only to the pronounced reduction of leaf nusber in the dry season. ‘They also enjoy an excellent protection by tightly closing stomate and alnost air-tight, strongly reinforced epidems. It seems difficult to avoid the conclusion that the maximal reduction of transpiration _restricting water loss in Larrea divaricata to 1/78 and in L.cuneifolia evento 1/150,1s at least partially due to the varnish whose layers reach in the latter speciés a thickness of 25 to }0 microns. Such extrene ratios of maximm to minimum transpiration (Seybold, 197) convincingly denonstrate the usefulness of the various layers tightening the epidermis of xeromorphous leaves in times of stress. This conclusion is also underlined by Morello who states (138,11; p.489) "It seems to remain finally proven that the structural peculiarities of the clerophyll- ous plants with low cuticular transpiration are responsible for this spectacular restriction of the transpiration current’ Adsorption of ethereal oils The resistance to the escape of water vapour fron the leaves can also be enhanced by a film of volatile oils excreted by glands, in analogy to resins. Recent work by Atay (10) corroborates earlier results ty Heilbronn (64) demonstrating that such films of oils fron thyme, peppermint, eucalypt, etc. appreciably reduce (a) evaporation of water from open vessels; (b) total transpire- tion of non-odourous leaves, if adsorbed fro the air, Heilbronn (65) vas further able to show that the rise of water in capillary glass tubes is considerobly reduced by surface adsorption of ethereal oils from a distance of 11 um. They can thus be expected to produce a drying effect on microcapillary systems in cell. valls if water colums recede from the surface thus reducing also the steepness of the gradient of weter vapour pressure. Some reduction in capillary rise could also be produced if, instead of blotting paper saturated by a drop of ofl, living leaves of marjoram, peppermint. or flowers of Jasmine functioned as. donors of oil. in a closed atmosphere. In order to obtain more spectacular effects it was necessary, however, to crush the leaves; they often would not release any oil vapour without éuch human help, as for instance the leaf of Mentha silvestris ‘brought in winter from outdoors into a heated laboratory. re Further studies under dry and hot conditions will be necessary to clarify definitely the problem of the ecological significance of volatile oil in plants. The present state of knowledge shows that much of the criticism of the naive period of plant etology. by Audus and Cheethem (11) and others was certainly justi- fied. The oils neither reduce leaf temperature appreciably by absorption of heat energy in the adjacent atmosphere into which they evaporate, nor is their volatility and their consequent cooling effect as great as was expected. But on the other hand Heilbronn's and Atay's experiments certainly have shown that ‘the effects on transpiration of adsorbed films of these, substances deserve eamest further attention. Future studies ought to distinguish clearly between effects on stomatal and cuticular transpiration since regulation by stomatal closure, little studied by Heilbronn but found important by the British authors and by Hafez (58) might prove decisive. ws/039.h2 *¥8/AZ/"15 - page 38 A pathological increase of water permeability of the cytoplasm in cells exposed to the vapour of volatile oils has been established by Atay, while effects on cell meubranes might be complex differing probably in cutinized walls and in cellulose menbranes of mesophyll cells with a more open microstructure. Finally ve wish to mention here tvo recent Turkish contributions to the Imdwledge of xeromorphous leaves and other organs.’ “Aykin (15) shows that stomata sunken in depressions or crypts - or else on epiderms protected by hairs - possess often pronounced hygromorphous properties: This is in accordance with their development in relatively moist air surrounding them during the process of thei development, On the other hand we learn from the investigation of Sayi (191) on Onobrychis ari that the silky-hairs covering the surface of the leaves in ary weathér, False themselves in moist air, thanks to a joint-like, hygroscopic mechanism of their bosal cells. Tt would seem that hereby transpiration is restricted in the first, but facilitated in the second case. ‘VIZ. DESICCATION RESISTANCE OF PIANTS IN DRY AND HOT REGIONS 1. General survey of concepts While resistance to drought of plants as a whole is a complex constitutional property (see Prof. Stocker's definition of “constitutional drought resistance” in chapter 3), the resistance of the organs, tissues and cells to dehydration called "desiccation resistance” 1s one of its fundamental components. In our gpinion, it depends ‘neinly upon the specific biochemical and physico-chemical constitution of the protoplasm of which little is known though certainly there are.also mechanisms of menbrane structure favouring desiccation resistance. Desiccation resistance is relatively easy to define and to measure. We can determine the water content of an organ, such as root or leaf, at saturation on the one hand, and at verious degrees of dehydration on the other - which in ‘turn might be harmless, dangerous or destructive. The water contents thus established can be simply expressed as fractions or percentages of fresh weigtit: ineluding vater, or of dry weight, determined at 105°C. in an oven. On the other hand, we canelinfnate dry weight - which changes with age and hour of the day - and express the resulting figures of water content as percentage of water content at saturation: this then is the "relative turgidity” of Weatherley (243). If, alternatively, we prefer not to insist on the water content remaining at each stage, but on the complement lacking to saturation, we are led to the concept of "water saturation deficit" (WSD) which is, according to Stocker, the difference detween a given water content Woot, and the maximum water content Wx expressed as percentage of the latter: : wsp = “nox 7 "nat x 100 Me In co far as the damge caused by desiccation 1s concerned, we distinguish "sublethal" (Oppenheimer, 147), “lethal” and "critical" (Hifler, 7h; Pisek and Berger, 165) vater saturation deficits, or else "critical", “lethal” (Marshall 120) and ‘bublethal" (Pisek and Berger, 168) water contents. These concepts have been variously defined and the reader is referred to chapter 3 of the present publication and to Levitt's recent took on "Hardiness in plents*. W8/039.42.WS/AZ/¥15 - page 39 On the other hand the reduction in water content will be almost invariably accompanied by a rise in the "osmotic potential” (Oppenheimer, 145) or, meaning tne same, “cemotie pressure, value, aiffusion pressure deficit” of the cell fluids. ' Therefore, many authors: following Walter, study the rise of this indicator of water strecs from normal figures (which are, as a rule, genetically fixed) to euperoptiml ones. The highest endurable figure found when irreparable dames just sets in, has been defined by Walter (242) as maximum osmotic value Onease)* ‘nae For the purpose:of the present review we do not discuss elther changes in osmotic potential or such in suction tension of cells and tissues called also their diffusion pressure deficits, in contrast to those of the cell fluids per se. Instead, we refer to desiccation recistant plants in terms of decreasing water contents or increasing saturation deficits since relatively more data have been accumulated on these by ecologists, horticulturists, agronomists or foresters even if little acquainted vith the concepts and methods of modern water physiology. In studies undertaken in the last thirty years, the authors sometimes dis- tinguish between water easily evaporating from a plant, the so-called “rree" water, and that fraction vhich is tenaciously held back by the colloids and solutes of the cell, the so-called "vound" water. This distinction, although disputed ‘by Weismann (246) is fundamental and paralleled by similar, (though also not clear- cut) classifications of soil moisture - above and below the "wilting percentage" or “hygroscopicity" of the soil. Recent but as yet wmpublished research by Dr. J. Cuayen in London suggests that some water in desiccating cells might be protected from evaporation being incorporated in certain lipoproteins of the protoplasn ‘hat Dehave as hydrophcbous bodies. In the latter state, hydrophilous groups binding water are supposed to be tumed inwards thus enjoying protection by the hydropliobous groupe forming the outer layers or shell of the Macromolecules. These lipoproteins may be of a semi-cristalline character, retaining vater in a manner similar to crystals of certain salts. Sone plant tissues are killed by even very slight water deficits. Such is ‘the behaviour of many hygrophytes. Other plants survive a high WSD which might reach 60, 70 or even 60 per cent ani more of thelr saturation water, Here the specific resistance of the protoplasm to desiccation is high, and we may expect. that, at least in certain cases, some bound water is kept back as an "iron reserve". Denaturation of the protoplasmatic colloids seems to be avoided by its presence. Finally thore are many cases. in cryptogms, but rather exceptional ones in highe= plants, where the cytoplam can becone air - and even oven-ary without undergoing fatal denaturation with practically no water remaining. We call this group “resurrection plants". Such plant types including many lichens, mosses, liverworts, fers and selaginellas can be moistened and dried almost indefinitely. They seem ‘to have attained the highest degree of perfection in their adaptation to hard terrestrial and epiphytic conditions. “We find them on very shallow soiis, in rocky depressions vhere vater accumulates for a short time after rains, and even on bare rocks, walls, etc., heated and dehydrated by the sun. They switch over from latent to active life within 2h or 48 hours. Their well known plasuatic desiccation resistance rms through an enormous gamit from, so to speak, zero to infinite-absolute. Thus, vhile thé phyeico-chemical mechanisms are little known, it is-not surprising that certain plants of aria rather than other. countries possess such water-retaining mechanisms, We shall discuss here grasses, conifers, epiphytes, resurrection plants and others. w5/039.41,5 /Az/415 - page Ho 2. Grasses ‘The extreme drought resistance of wheat, rye and barley tn very early stages of germination has already been observed in 1827 by de Saussure (190, see also Levitt, 107). A modern study on wheat was recently published by Milthorpe (131) who, like Saussure, used concentrated sulphuric acid to dry the air in low humidity chambers where he tested seedlings at consecittive stages of germination. He was able to show that three days old seedlings with coleoptiles not longer than 5-4 mm could survive a loss of 98 to 99 per cent of their totel water content. The shoot of older seedlings, up to 17 days old survived a 90:per cent WED while roots longer than 2 mm vere killed by an 80 per cent or less WSD. Undeveloped root initials, hovéver, can stand ® WSD of 96 per cent,-& clireumstance highly important for the survival of cereal seedlings in semi-arid comtries where a prolonged ary Period often follows the first rains of the season. Though the first two leaves, if already developed, will be damaged already by @ WSD of 30 per cent, the vegeta- tion point of the shoot will replace then. Desiccation resistence in wheat is a varietal property. As Bayles, Taylor and Bartel (16) demonstrated by eradication of field grown plants before the “pooting" stage, some varieties will part with their vater more readily than others. Hop, a variety sensitive to drought, retained after 2h hours only a water content of 52 per cent per fresh weight, instead of the original 60. Marquis, intermediate in its behaviour, kept 57, but the drought resistant Kubanka of South-Russian steppic origin, as much as 63. Whiteside (254) studying reaction to’ soil drought, found an important increase of desiccation resistance. While parenchynatic leaf sheath tissue of plants grown at high sot] moisture contents survived in drought chambers above 0.6 mol common salt solution, hardened tissue was found alive above 1.4 mol and same cells even withstood 2.6 mol meaning about 150 atmospheres of suction tension. A surprisingly high desiccation resistance of an annual grass, was established by Killian (94) at Béni Ouif, in the Sahara. Its leaves will not dxy up even if only 30 per cent of water per fresh (42.8 per dry) Weight, remain snd their roots are even more resistant to desiccation. These properties, togéther with the activities of very long but thin roots, ensble this Peamophyte to withstand reinless periods, as well in the seedling as in the fruiting stage. As the author points cut, it would be quite erroneous to, consider all annuals growing in deserts as drought escaping ephemeres, 1,e. neither needing nor possessing xerophytic adaptations. Perennial grasses of seitt-arid pastures resenble wheat seedlings in their ehaviour to drought. They will appear dead during the dry season but, as Walter points out (2k1) the youngest leaves of their shoots, the leaf bases and the well protected apical meristems reuain alive. Even some adult leaves looking brom, ‘as is the case in some species of Aristida studied by Henrici (66; p.62h) and in contortus seedlings studied by Giendening (52) in Arizona, will regain a Goch SolGUFe: Internal stele tesues of stotona and voote ofven reaain alive, in spite of drought periods lasting for many months. They even preserve a, considerable water content: Walter found about 70 to 90 per cent water in tiny young leaves, Thus desiccation resistance of these grasses is rarely clearly apparent, except in the seedling stage. ater, the plants give up the majority of their mature stems, only inner strands in stolons and roots (Henrici, 66), more Ws/039.41WS/AZ/415 - page 41 or less embryonic initials of shoots, leaves and roots remaining alive. These often resume growth after @ surprisingly short interval subsequent to the first reins, Glendening saw some “greening” after pronounced dormancy in Heteropogon on the day following the first rain and rapid grovth a day later. Miss Henrici (66,11; p.€23) writes on the grasses of the Armoedsvlakte after vain: "Grasses developed with incredible speed by shooting from the margins of old tufts but not as individual young plants", She found fev grasses restricting their transpiration during times of physiological activity. This, of course, produced considerable saturation deficits of the leaves corresponding often to a loss of 20 per cent of fresh weight, but sometimes even of 50 per cent without causing damage. It was remarkable that high deficits were often not. connected with drooping of the leaves. The latter seemed to preserve high turgidity ‘thanks to # high capacity of elastic contraction of their cell walls. If dehydration was temporary, this mecbaniem seemed to be highly effective in maintaining life; however if permanent wilting occurred the leaves dried up within a few days. However, the resistance of grasses to dehydration in the grasslands of the world evidently varies widely according to species. Mueller and Weaver (139) experimenting with Bouteloua gracilis and hirsute and with Buchloe dactyloides Lypleal cpecten of the shore grass pralzis of Nebrasia ~ Pound those species extremely resistant to dry and hot air. When air at 64°C. was blown on them during five hours, they not only preserved their vegetation points but even most of their adult leaves. This proved possible, however, only if water suyply from the soil continued; even these grasses will dry up if this ceases, their leaves Deing unable to withstand prolonged wilting. Whitman (255) investigating the resistance to drought of grama grass, Bouteloua gracilis, and two other xeric grasses, vas able to show that under ary conditions, Bouteloua spends 96.5 per cent of its free water, retaining only the ‘bound water Which vepresented not more than 37 per cent of dry substance. While ‘this grass is unable to increase its bound water during drought, the preservation of life at such a low water content is exceptional in flowering plants. Quite often ve find leaves even of pronounced xerophytes drying up if their water content Decomes approximately equal to the dry matter, Whitman concludes that bound water changing little during exposure to drought offers no indication of drought resis- tance. Its rising share in total water after such periods did not, in the plants studied by him, reflect active increase of bound water by synthesis of water binding colloids. It merely reflected the dryness of the habitat. The author admits, ou the other hand, that a highly drought resistant species might possess a high proportion of bound water as a constitutional character, as is actually the case with certain desert plants, studied by Migahid (128). ‘The wide variation in resistance to dehydration.in grasses is very obvious in the results of Paltridge and Mair (161) in Australia, Experimenting + aoe long array of pasture grasses, like Brac (ium, Lolium, s sy they tected their vater fing properties after eradication from the Boll, After the initial heavy losses by transpiration the species entered a phase of low losses in weight accompanied by permanent wilting. This phase set in at different levels of water. content ani lasted for quite different periods, before ‘the plant finally dried up. ¥5/039.4.WS/AZ/¥15 ~ page 42 Choosing the pronounced drop in transpiration at the beginning of obvious wilting as 8 criterion of drought resistance, the authors classify species which at this moment preserve 75 per cent at least of their water at full turgidity as true mesophytes. - Between 50 and 75 per cent the species are called xerophytic mesaphytes; between 25 and 50 per cent mesophytic xerophytes; below 25 per cent true xerophytes. The definitions mde in the spirit of Meximov prove of prectical ‘value and useful for a first orientation corresponding on the whole to the ecolo- Glcal character and distribution area of the species. Thus, Themeda australis Wilting just above the limit set for mesophytes, 1s a species groving in forests with more than 30 inches of rein but penetrating into drier regions. On the other hand is mildacea wilting only at 10 per cent (1) and Stipa nitida at 14-20 per cent relative turgidity (when we should certainly expect thar tot been completely dry and dead), grow under rether dry conditions, the first in the mediterranean area, the second in Australia, Appreciating theeffort of the authors to find an objective measure of xerophytian by resistance to desiccation and preservation of life during permanent ‘wilting, we doubt its general applicability since constitutional drought resistance is far more complex than dehydration resistance. Thus Henrici's studies vith grasses led this fine physiologist to a different interpretation of xerophytism. Although admitting thet restriction of transpiration is an important xerophytic character, she would not concede that the capacity to withstand prolonged wilting represents a valid criterion for the grasses studied by her. Xerophytiem is far too complex to be adequately defined in so simple a manner and it is little surprising that Bailey (14) found no correspondence betveen the critical water deficits of three Anerican prairie species, as defined by Paltridge and Mair, and their drought resistence. 3. Conifers Generally speaking the desiccation resistance of conifers of hot and dry regions is less extreme than might be excected if we consider their penetration into regions with prolonged dry periods end the pronounced capacity of certain [pines and cypresses to restrict their transpiration (Henrici, 70). Marchali (120) defined as “critical” water content of conifer seedlings the fraction remmining over when leafy organs change colour from dark to light green and hypocotyls or cotyledons begin to appear wrinkled. He found these symptoms of damage in seedlings of westem species at the following figures of relative turgidity: Tmja occidentalis - 0.85; Abies is - 0.75; Pinus rose, monticola, Saribses - 0.79, 0-64 and 0.50 Sec Seaetraly; 5 Mauge hater = 02 Tmja Ticata - 0.28, No clear trend towards lower criticsl vater contents with’ habitat can be derived from these figures. Parker (162) worked with species of the seme region in Idaho but tested the needles (or, in Thuja, the shoots) of mature specimens in winter when desiccation resistance reacbes its highest peak (Pisek and larcher,170). - He was able to preserve needles in a Living condition with only 60 per cent water calcwated upon ary weight in western yellow pine and with 68 per cent in Douglas fir while the ‘tranchlets of Thuje ta, remained alive with mich’ Jess, water, in agreement with Marshall's W5/039.42.w8/az/¥15 - page 43 What matters from an ecological point of view, is in Parker's opinion, the ‘time elapsing in isolated organs, before death occurs, rather than the lethal level of water loss.. It seens therefore importent that the leaves of western yellow pine (Pinus ) could be kept living for 12 days at 40 per cent humidity and 3e of Douglas fir lived under the same conditions for six days. Here lies the advantage over Pinus nonticola, Abies is and Picea mannit and also licata whose needles éry up mic = In another paper ‘the author vas able to show that one year old needles of Pinus lexose keep living in the laboratory for'tvo to three weeks vhile Older ones HGEGRSE one’ or tor, In these experinents the needles dried wp-atter having Jost about 50 per cent of their original total weight. Their desiccation resistance vas very much higher than that of Pinus strobus and Penigra investigated at Mich (Parker, 163). These began already to suffer after & loss of only 10-15 per of their original veight vhile death took place after a loss of 25. ° Pinus cenbra and Picea excelea studied by Pisek and Larcher (170) in the Alps show @ behaviour very similar to that of Western Yellow pine and Douglas fir in the Rocky Mountains. At the winter peak of their resistivity to desiccation, their needles withstand dehydration till a point where only 55-70 per cent of the water related to dry weight remains. Such resistance appears rather extreme if the bigh proportion of thickened.cell wells in conifer needles is considered. But Pinus silvestris is, according to the authors, even more resistant to desiccation. At the stb-lethal level the water content of its needles can drop ‘to less than 40 per cent of the dry weight, . If we compare these figures vith those of the Aleppo pine whose needles will suffer scme damage if their water content ‘drops below 100 per cent of ary weight (unpublished results of the present author) we meet with a remarkable discrepancy Between resistance to desiccation and physiological drought resistance which also comes forth if we compare the possible duration of exposure of isolated needles or brachyblasts to dry air. Needles of Pinus silvestris, a tree of the Iumid and cool zone, will keep alive for seven days (Pisek and Winkler, 169), those of the Aleppo pine (except 4f remaining commected with the isolated mother branch) less than two, under sumer conditions in Isreel, ‘The solution of the discrepancy seems to be thet northern conifers might be subjected to heavier vater stress during the frost periods of the cold winters of their habitats, than possibly mediterranean pines even during sumer when some water will still be supplied by the roots and stomata will not be fomd hermetically closed but remein open if we pluck needles on normal sumer days. Authors like Schormeyer (194): and Parker (163) have investigated pines in the expectation that.the more drought-resistant species should also possess the higher desiccation resistance. Schoymeyer found, however, in Pinus echinats, considered as more drougkt-resistant than P.taeda, neither a greater quantity of bound water at the permanent vilting percentage nor a higher omotic potential. The min reason for the higher drought resistance: seemed to be a better developed root system, a situation which proved also decisive in pines of the Great lakes’ region (Shirley and Meuli, 200). Parker compared needies from the relatively mesophytic Atlantic-continental White pine (P.strobus) vith that of the slightly xerophytic P.nigra. Their water retaining pover (rate of loss in weight when kept isolated ‘water) ran contrary to expectation. It was higher with P.strobus, and there was practically ho difference in the critical and lethal watgr contents. W8/039.41‘WB/AZ/415 - page bh There is more than one to the understanding of these disappointing results: (1) Ag pointed out by Stocker (218), desiccation resistance is by no means a necessary element of constitutional drought resistance. (2) lio correlation between desiccation resistence and ecological-geogrephical distribution can be expected if only sumer drought is képt in mind. Winter and frost resistance as a potent factor in distribution and resistance should consequently always be considered also, Actually the White pine might be exposed in the eastern states of the United States of America to heavier and more prolonged atmospheric and soil frost than at least the mediterranean eco-subspecies of P.nigra, and this might prove decisive for the more pronounced water retaining capacities of the former species. Generelly speaking, we think that conifers are a very heterogeneous group, phylo- genetically and ecologically, containing various "hydroeconomic" types (Zchary, 260) e6 much as other evergreens: There may be considereble differences in behaviour to frost and drought even in ecotypes of the same species, While a discussion on constitutional drought resistance is beyond the scope of the present chapter, the following recent results with forest trees ate:mentioned in passing: (257) grev seedlings of pines, cypresses and other trees in containers. After ‘reduction of soil moisture to the permanent wilting petcentage, their @rought chamber. Agreement with field behaviour wes incomplete. Pinus brutia poral es drought-enduring than P.I end Eucalyptus camaldulensis rostrate) more than ‘gemphocephala, as Weald be expected on inoue resistance in jeries and plantations. ‘Supresees: sempervirens, however; Proved much more Se ee alten te mediterranean conifers persisted Ito jy) the eycalypts and Casuarine torulosa for 4 to 6, while Tamarix to draw much water even in sumer from desertic sand dimes, died 3 days in spite of the fact that 1t occurs naturally in the driest region. contrast to the tested conifers, it is not a xerophyte in the sense of Maximov. 4, Mediterranean sclerophylls We find similar differences 2180 between broad leaved mediterranean Selerophylls. While Oppenheimer’s (146) and Rouschal's (18%) early investigations on species of oak, carob, stone-linden (1 ), laurel, ete. rendered on the whole sublethal WSD's between 60 and 70 per cent, there are cases of much lower desiccation resistance, e.g. that of Pistacia lentisous. This shrub is e pioneer on eroded and strongly insolated rocky sites and dederves to be called highly eat On the other hand, leaves of the Isurel, decidedly Iess dtoudit-resistant consti- tutionally than carob and oaks (Boyko, 23) will keep on mch loriger without water supply than the leaves’ of the latter and suffer.a higher WED, of 60-65 per cent, without damge (the author's ovn recent investigations). The species whose leaves possegs a highly viscous protoplasm, resembles in this respect’ the olive tree. .Carobs and oaks keep a medium position in this respect between the highly resistant Inurel-olive group and the very sensitive lentisk. Carcb leaves evince sate dry spots already if their vater coutent drops below their dry weigit. They require a higher water content than the leaves of ilex which have equal Gry weight and veter at saturation tut will be able to 19 per cent of their saturation velght (=50 per cent of maximm water content) without suffering serious damage. seer Voureocetsene) gelliprinos, much resembling the kermes oak (Qu.coccifera) of the west WS/039.41WS/AZ/¥15 - page 45 mediterranean basin, absorb at saturation up to 60 per cent water of their total veigrt. Of this water 2/5 can be lost without serious damage, i.e. their sublethal water deficit is 40 per cent. Thus both species are considerebly more desiccation- resistant than the carob. Leaves of this tree keep, at the dangerous sublethal level, only about 10 per cent water per fresh weight corresponding to about 70 per cent of ary veight. ‘This figure is, however, still far above that of the laurel leaf vhich will be found still intact with a rest of water about 50 per cent of its dry weight (Oppenheimer, unpublished work). Indeed, we found in October a sublethal water content of 44-49 per cent of ary weight in Laurus nobilis the mature leaves of vhich contain less vater than dry substance even at saturation. “Comparing meditérranean evergreens with highly desiccation-resistant dvarft shrubs, we find that laurel and olive reach the same degree of desiccation . resistance as sordidum studied by Meier (12h). This rock plant growing on. ary and barron atopes “scorched by the sun in southern France, was found to have apparently living leaves at little more than 50 per cent water per dry weight. Both evergreens further approach, by their extreme desiccation resistance, the evergreen alpine ericaceous dvarf bushes Arctostaphylos, Rhododendron, Vaccinium vitis-ideea studied by Pisek and Winkler the first mentioned is able to remain in life with only 30 per cent of water per dry leaf weight! Our present investigations on the desiccation resistance of mediterranean sclerophyll leaves aim among others at an improvement of the diagnostic principles used for the determination of damage caused by drought. A leaf is not only considered as damaged if dry spots appear on the epidermis or in the mesophyll ‘but also if it is found unable to return to its initial saturation weight if offered the opportunity, in correspondence vith the definition of a limiting value (Grenzwert) by Arvidsson (6; p.24), Further serious attention is paid to the iscoloration of the bundle sheaths of the min ‘or secondary nerves where death otten occurs first - probably because the cells are relatively bighly hydrated and vacuolized. 5. Other ecological groups The Literature on desiccation resistance of flovering plants is still rather scanty and some older investigations lack a modern outlook. Among the older works we shall mention the study which Kamerling (84) carried out in Java and Brazil. He found epiphytes like Dendrobium secundum and Sophronites cera highly resistant to desiccation. “The former when kept without roots and water supply in the shade, lost only about 0.35 per cent of ite fresh weight per day. ‘Thus it kept.on for 62 days without any loss of leaves worth mentioning, A spécies of Tillandsia, a heliophyte from the surroundings of Rio de Janeiro, lost in 32 days only 17.25 per cent of ite fresh weight and finally only 0.4 per cent per day. salis cos: ip agreenient with other non epiphytic cacti was cat ae te Sete able to Fration to 0.15 per cent of ite fresh weight per day, losing finally 65.5 per cent but suffering damage. 1m vaccinifolium @ fem losing at the beginning 4.7 per cent of its we! at y only 0.27 per cent but at the final point (71.2 per cent loss) its leaves had dried up. W8/039.42.ws/nopnas = page 46 ‘the plants stuiied which the author considered as the real and true "xerophytes" are either succulents or, as Po! ium, near relatives of tropophytic ferns suffering complete desiccation, Their ‘water losses are in many cases connected with xeromorphous structure, like thick cuticles, water stoving tissues and water absorbing hairs. The water balance of these epiphytes is ectremely interesting biologically but of limited practical importance. The interested ‘Turning back to terrestrial xerophytes, we find very interesting indications in @ stuty by Thoday (222) on Passerina filiformis groving in the Cape Province. The needle-like leaves of this plant bear stomata only in a groove on their upper side which 1s protected at its margins by hairs and closes at low humidities. ‘Thoday found the branchlets of this plant to have even at saturation only 47.2- of water in their fresh weight - nearly as the above mentioned Quercus Tex. The grooves close up at 3% per cent but the leaves were still found ‘Tiving at 29. This figure meaning that the branchlets preserved only 41.4 per 6. Resurrection plants Studies carried out on plants suffering desiccation to air dryness seen very -promising for the understanding of desiccation resistance. Leaving eside mosses and lichens, relatively little has been achieved in the physiological study of this remarkable biological group containing spermatophytes such as Careyphysodes growing in Transcaspian sandy deserts (Vassiliev, 251), Ramonda nathaliae and Myrothammus flebellifolia ~ cf, Miller (130) - rock plants growing, the former in ‘the Balkans, the latter in Rhodesia. There e,clse Chamacel ene snbrepioue fron South-West Africa, resenbling a water lily in the vet season to air dryness in the rainless months. A few botanists have atudied revivescent ferns.We shall mention here the work of Pessin on Po um ides (167). The author found little water retention by the fre: ap sphere-and concluded that the scales covering the lower flank of the fronds serve water absorption by the dry plant. After a week in an exsiccator the plants still preserved 25 per cent of their original water content. In contrast to this, Rawitscher, Hueck, Morello and Paffen (180) established that uprcoted plants of Sei convolute groving tn ‘the Cantinga in Brazil kept, in an air-dry condition not more =10 per cent of their saturation vater content - meaning 14-18 per cent of dry weight. The eubling in its biological properties the better knovn Se idophyla from Central America, absorbs much water externally ‘periphery, we [on excluding in most cases cuticular retention. 88/059.41.W8/AZ/M15 - page 47 We shall also mention a study by Rouschal (185) on Ceterach officinarum. This little fern, widespread in mediterranean and adjoining European countries, also lacks water-retaining mechaniens. It loses water efter a rain, like a sponge (up to 147 per cent of its dry weight per hour). Finally 10 per cent of the saturation water is retained by physical forces. But healthy and full. grown fronds will withstand even desiccation in a closed chamber over sulphuric acid and return to life by rehydration absorbing Liquid water or vapour. Repeating Rouschal's experiments we were able to corroborate these findings and also his assertion that the vacuoles of the drying cells become solid during dehydration ~ @ process earlier observed by Iljin in the similar fern Nothochlaens marantee (80). This together with collenchymatous thickenings of the call reduces far-reaching distortion of the tissues in an air-dry state, Working with this revarkable plant, it is highly interesting to convince oneself that epidermis and chlorenchyma cells return indeed from an anabiotic shrunken to a completely normal condition in the course of about 36 hours. They can be plasmolysed and emit fluorescent light - in contrast to dead cells of the same species. Their plasm locks completely homogeneous, while we found it reticulate in the ary epidermis cells of the peduncle vhich had been evacuated before in liquid paraffin fron air hindering observation. ‘The meshes of the ectoplasm measured about 2 or 2 microns across (Oppenheimer, unpublished work), The plasm forms in its ansbiotic state flat ribbons and tongues lining the walls and apparently also spread out across the cells. Net - or rather honeyconis-like structures in the ectoplasm or the adjoining walls of desiccation-resistant plants have been observed by various authors: ‘by Renner (185) in Eriopus remotifolius, by Hitler (7h) in liverworts of the Jungermemiacene family and by Simonovitch and Levitt (207) in dehardened cortical celis of Hydrangea. In the latter case it has been interpreted as possibly produced by torn cytoplacmatic strands of an easily dehydrated ectoplasm. While this sounds quite plausible, the phenomenon might also point to other reticular structures such as ribonucleic acids, etc, which might become visible ea upon dehydration. Their connexion with desiccation resistance is recommended x study. ‘7. Theoretical foundations of desiccation resistance Concluding this review, we wish to express the conviction that future research will synthesize the now conflicting theories regarding desiccation resistance. As pointed out by Stocker, (see chapter 3), Iljin argued that death of the plasm during cell desiccation 1s exclusively due to mechanical stress. In his opinion the cytoplasm is apt to be killed if exposed to tension during the evaporation of @ central vacuole, compressed by shearing forces during shrinking or forcibly extended vhen to its original dimensions after pseudoplasmolysis. As pointed out by Levitt (107) I1jin's theory agrees with most Imown facts regarding @esiccation resistance. We aiimit that his reasoning about the influence of cell size and shape on the degree of tension seems physically sound: a small cell possessing a relatively large surface will suffer less tension per surface mit during dchytretion than a larger one. But this principle must not, of necessity represent ‘the key to the explanation of small cell size in xerophytes. The euall cell size might more simply and conveniently be interpreted as a consequence of phylo~ and ontogenetically effective scarcity of water during develomment. And T1jin's conviction that since of two cells of equal volume that possessing the more developed surface will be under less mechanical stress and that consequently the ws/039.42NS/AZ/"15 - page 4B spherical shape is the most unfavourable one for desiccation resistance, is certainly not borne out by the actual structure of most drought resistant plants, Isodismetric cells are the rule rather than the exception in xeromorphous plants. In @trect contrast to T1jin's principle, already Lothelicr (112) was able to show that under ary conditions epidermis cells of dicots became uct less but more isodismetric. If, on the other hand, palisade tissue developing in ary surroundings is found camposed of narrow and elongated cells - in spparent agreement with I1jin's expectation - this is as meitioned above, probably rather an adaptation to atrong light, instead of drought. Considering the tearing apart of cytoplasm during dehydration, it should bé recalled that this phenomenon called "plasmoschisis” merges into plasmolysis which is not necessarily harmful. It should be further recalled that very often one observes the separation of plant protoplasm into several portions, without fatal consequences. If the foretble separation of the bulk of the protoplasm from the wall end that part of ‘the ectoplasn intimately comected vith it, is fatal in some cases but harmless in others the explanation mist be looked for in differences of elastic properties studied by Scarth and his associates (192,207). Differences of molecular constitution end subnicroscopic structure might be expected to be more decisive then tensile strength. That tensile stress 4s in itself dangercus to protoplasm even if not followed by plasmoschisis has not been demonstrated by Ijin. His ‘theory, though useful for the explanation of certain phenomena suffers, in our opinion, of an oversimplification of the facts and does not cover all phenonena, In particular we feel mable to accept I1jin's assertion that the vegetable protoplasm, irrespective of species and organ, withstands complete desiccation Provided that rehydration is carried out gradually and carefully. Levitt (107; P.1h9) speaking in this connexion of “incompletely killed" cells, is probably quite right. We doubt if more than the tonoplest survived in the epidermis Slices of Rhoeo discolor or red cabbage which bad been kept over concentrated sulphuric acid fn Tijin’s experiments... Thirty years of fruitful research on “protoplasmatics” at Austrian universities by Weber, HBfler, Biebl and others has produced a vast bulk of information regarding specific structural differences in the protoplasm of systematically and ecologically different plant grows. That specific structure - of vhich so far relatively little is nom - seems to play an important part in cases where desiccation resistance is not explainble by Plasmoschisis, has been shown by Héfler's (74) studies with liverworts and his subsequent publications. Abel (1) continuing these studies with mosses arrives at the conclusion that desiccation resistance in mosses varies with age, season, and temperature and is essentially a consequence of specific protoplasmic properties, at least under field conditions. Mechanical damage plays a more modest part mainly in lsboratory experiments if changes in water saturation are very abrupt. With mosses and livervorts, a classification into xero-, meso- and hygrophytes is possible on the basis of desiccation resistance alone. But, as we have emphasized, this seems impossible with spermatophytes where constitutional drought resistance, decisive for classification, bears a more complex character. w9/039.41.8/az/"15 - page 49 Regarding the so far hypothetical specific plaamatic properties we dare not add much to what bas been already said above regarding lipoproteins, The higher density and ductility of resistant plasm will have to find its place in. an embracing theory of resistance for which Bogen (21) and Stocker (218) heve contributed building stones. Possibly the most essential task is the explenation of how to avoid the denaturation and precipitation of constituent proteins. Molisch (133; p.97) has discussed this as well as the precipitation preventing substances, the possible differences in the components of proteias such as peptones and of alkalinity, etc. in this comexion, Recent experience of Kessler (90) further hints at e direct or indirect influence of nucleoproteins on drought resistance of higher plants. WS/039.41