Interacciones Planta‐Nodriza en Lophophora diffusa (Cactaceae) en un Desierto Subtropical de México1

One of the two species from the genus Lophophora is the false peyote Lophophora diffusa, an endemic cactus species of the xerophytic shrubland at Querétaro, México, considered threatened from illegal extraction due to its hallucinogenic and medicinal properties. We analyzed the spatial distribution of L. diffusa and its association with the locally dominant shrub species with the system SADIE (Spatial Analysis by Distance IndicEs). We also studied the principal microclimatic factors (light, temperature, and humidity) beneath the canopy of the possible nurse plant species. The use of SADIE is a new approach to study the mechanisms of spatial distribution. Lophophora diffusa and the shrub species presented an aggregated distribution with patches and gaps, as indicated by the values of I_a= 4.179 for L. diffusa and I_a= 1.660 for the vegetation. Lophophora diffusa was positively associated with the arboreal vegetation, particularly with Larrea tridentata and Acacia sororia, but was negatively associated with Celtis pallida and Myrtillocactus geometrizans. Microclimate evaluation indicated that C. pallida canopy significantly reduced radiation and temperature compared to the other species (L. tridentata, A.sororia y P. laevigata). We had expected L. diffusa to exhibit a positive spatial association with C. pallida; however, the reduction in light availability apparently limited seedling establishment of L. diffusa.     

Isozyme patterns in callus cultures and in plants regenerated from calli ofCereus peruvianus (Cactaceae)

Electrophoretic patterns for isocitrate dehydrogenase (IDH; EC 1.1.1.42), acid phosphatase (ACP; EC 3.1.3.2), peroxidase (PER; EC 1.11.1.7), and esterase (EST; EC 3.1.1.1) isozymes were determined inCereus peruvianus tissues and used as markers of genetic uniformity of calli and of the plants regenerated from callus cultures. One IDH, six ACP, six PER, and six EST isozymes were induced in cultured callus tissues in medium containing three 2,4-dichlorophenoxyacetic acid and kinetin combinations. Four ACP, two PER, and three EST isozymes were still present in all regenerated plantsin vitro and therefore can be used as markers of theC. peruvianus plants regenerated from callus tissues. The differential patterns of ACP and IDH isozymes and the similar zymograms for PER and EST isozymes presented by callus tissues were used in a comparison of callus tissues cultured for 2 years. The comparative analysis of zymograms within each enzyme system indicated a mean heterogeneity coefficient of 0.33 forC. peruvianus calli cultured for 2 years. Because of the isozyme variations, which developed in culture medium and were transferred to the regenerated plants, the IDH, ACP, PER, and EST enzyme systems can be considered to be good markers for investigating possible genetic variations in plant populations ofC. peruvianus obtainedin vitro from callus culture.This research was supported by the CNPq     

Testa sculpture in the genus Frailea (Cactaceae)

The primary sculpture of the seed surface in the genus Frailea (Cactaceae) is characterized by a i flat outer periclinal wall and the presence of an usually eccentric papilla. Papilla morphology differs mainly in size and breadth:length ratio. Seeds of different species (groups) are characterized by special patterns in the arrangement of short and long papillae on the surface, thus providing a useful tool for taxonomic studies in Frailea. The seeds of Frailea castanea possess branched papillae, a character restricted in this genus to this species. Moreover, such papilla morphology seems to be unique for the seed epidermis in spermatophytes. The possible function of the papillae is discussed.     

Application of randomization methods to study the association of Trichocereus pasacana (Cactaceae) with potential nurse plants

Spatial distribution of plants and seeds of Trichocereuspasacana was studied in relation to bush species abundance, and ofcacti in relation to the available number of seeds under bush species, in LosCardones National Park, Argentina. Randomization tests, carried out to find thesignificance level of Chi squared test statistics used to test fornon-random association hypothesis, proved that T.pasacana plants, seeds, and seedling establishment is related to thedifferent bush species that co-occur with it. Positive interactions werefound mostly with L. divaricata, but also withP.ferox and A. spartioides, suggesting that theyact as nurse plants. Negative interactions were found with B.boliviensis, A. spartioides, S.crassiramea, and Verbena sp, in spite of thehigh seed storage beneath their canopies. Cacti are more frequent than expectedin relation to seed relative abundance under L. divaricataand P. ferox and less frequent under P.rougesii, B. boilviensis,Verbena sp, G. glutinosa,S.crassiramea, and A. spartioides, due todifferential seed or seedling mortality when emergence occurs.     

Stem and wood allometric relationships in Cacteae (Cactaceae)

Allometric relationships in organisms are considered a universal phenomenon. A positive scaling has been reported between stem size and cellular size of tracheary elements in wood of different vascular plants, but few studies have been carried out in slow-growing succulent plants. The aim of this study was to evaluate if a relationship exists between size, growth form and wood cell size among individual species of Cacteae. Forty-four species belonging to 16 genera of the tribe Cacteae with differing growth forms and sizes were studied. When analyzing plant size, we found a positive allometric scaling and the larger-sized species showing a higher percentage of succulent tissue and less accumulation of wood tissue. The positive scaling found between plant size (height and diameter) and vessel elements and fiber length support the universality of the allometric relationship proposed for other vascular plants with non-succulent stems. Notably, wide-band tracheids do not scale with plant size or growth form. Succulence associated with narrow vessel elements with distinctive helical secondary walls and wide-band tracheids suggest they are the key adaptations to tolerate drought and provide support to the stems of most taxa in Cacteae. Fibers do not have the primary role of giving mechanical support; therefore, we assume the scarce fibers in clusters represent reaction wood that, along with the fundamental tissue, maintains the vertical position and shape of those species growing in rocky cracks. Our results with species having short succulent stems support the universal theory of positive allometric scaling of vascular plants.     

Non-concerted ITS evolution in Mammillaria (Cactaceae)

Molecular studies of 21 species of the large Cactaceae genus Mammillaria representing a variety of intrageneric taxonomic levels revealed a high degree of intra-individual polymorphism of the internal transcribed spacer region (ITS1, 5.8S rDNA, ITS2). Only a few of these ITS copies belong to apparently functional genes, whereas most are probably non-functional (pseudogenes). As a multiple gene family, the ITS region is subjected to concerted evolution. However, the high degree of intra-individual polymorphism of up to 36% in ITS1 and up to 35% in ITS2 suggests a non-concerted evolution of these loci in Mammillaria. Conserved angiosperm motifs of ITS1 and ITS2 were compared between genomic and cDNA ITS clones of Mammillaria. Some of these motifs (e.g., ITS1 motif 1, 'TGGT' within ITS2) in combination with the determination of GC-content, length comparisons of the spacers and ITS2 secondary structure (helices II and III) are helpful in the identification of pseudogene rDNA regions.     

Population variability in the genusTrichocereus (Cactaceae) in central Chile

Trichocereus has 4 species in central Chile. Population analyses indicate that introgressive hybridization has occurred betweenT. chilensis andT. litoralis in the Valparaíso and Santiago coast ranges. A taxonomic key is presented.     

Noteworthy idioblastic sclereids in the stems of Eulychnia (Cactaceae)

Stems of Eulychnia (a genus of six to nine species of candelabriform or arborescent cacti) have a parenchymatic cortex with two distinct regions. The outer chlorenchymatic layer is characterized by a conspicuous parallel striping, whereas the inner cortex region devoid of chlorophyll has a coarsely granular aspect. Stem samples from nine accessions, collected in the field or taken from cultivation, were studied from resin-embedded microtome sections and maceration. Two different forms of lignified sclereids were found dispersed in the cortex and the pith. The sclereids of the outer palisade-like cortex layer are distinctly elongated and strictly oriented at right angle to the stem surface, whereas those of the inner cortex and pith are globular or subglobular and conspicuously enlarged compared with the surrounding parenchyma cells. The ontogeny of the sclereids was studied from stem samples of different ages. Formation of the secondary cell walls starts only after cell growth is completed. A screening of numerous South American cacti for the presence of idioblastic sclereids showed that these structures are unique for the genus Eulychnia. Finally, functional aspects of the sclereids are shortly discussed. It is assumed that the sclereids contribute to the mechanical support and reinforcement of the plants.     

Chromosome number variation and polyploidy in the genus Echinocereus (Cactaceae)

Analyses of meiotic and mitotic chromosomes were undertaken in 16 taxa of Echinocereus belonging to 12 species and all seven taxonomic sections (sensu Taylor). Chromosome numbers are reported for the first time for eight taxa, and previously published chromosome counts are confirmed for the remaining eight. Both diploid and polyploid counts were obtained. Eleven (69%) of the taxa surveyed were diploid (2n = 22); the five varieties of E. engelmannii were polyploid (2n = 44). Overall, chromosome counts are available for 23 of the 48 proposed species (sensu Taylor). Of these, 19 (82%) are diploid, and four (18%) are polyploid. Polyploid cytotypes are most common in the primitive sections, e.g., sections Erecti and Triglochidiatus, which suggests that polyploidy is probably a derived condition in Echinocereus. Polyploid taxa range from medium to high latitudes and elevations relative to the overall distribution of the genus. Polyploidy, hybridization, and cryptic chromosomal rearrangements are thought to be the major causes of the speciation events of the genus.     

Physiology and genetics of self-incompatibility in Echinopsis chamaecereus (Cactaceae)

The self-incompatibility (SI) system of a geophytic cactus (Echinopsis chamaecereus Friedrich & G. Rowley) was examined in a series of experiments. Pollination tests indicated that E. chamaecereus is an obligate outbreeding species with a functional SI system. Incompatible matings were characterized by stylar inhibition of pollen tube growth and lack of fruit set. Two S₁ seedlings were recovered when plants of one clone were exposed to 42°C for 16 h and flowers were selfed immediately after incubation. The two S₁ seedlings and the parental (S₀) clone were crossed in a full diallel. Results were consistent with a one-locus, gametophytic SI system with two different S alleles. Disturbed segregation at isozyme locus Lap-1 was attributed to close linkage with the S locus (recombination frequency = 11±8%). This is the second report of close linkage between Lap-1 and S in the Cactaceae. Recevied: 1 February 2001 / Revision accepted: 13 March 2001     

Functional Morphology of Aztekium ritteri (Cactaceae)

The dwarf globular cactus Aztekium ritteri shows a high degree of surface differentiation. The stem consists of 8–15 ribs, these in turn being sculptured by transverse furrows: compared with a sphere of equal size, the surface/volume ratio is increased by more than 10-fold. At the same time, only 20% of the stem surface is exposed to direct sunlight, 80% lying in the shade of its own body, in a system of furrows and crypts. The exposed areas have few stomata and the epidermal cell walls are notably thickened and cutinised. In addition, these form an exoskeleton by means of a hypodermis containing calcium oxalate crystals while the unexposed cell surface regions, hidden in the furrows, lack xeromorphic characters and calcium oxalate crystals. By far the greatest number of stomata are found in the furrows mostly in deep crypts. When dehydration occurs as a consequence of longer periods of drought, the furrows and crypts close together, thereby reducing transpiration losses. The complicated interlocking nature of the surface architecture of Aztekium is discussed in terms of the functional aspects as an adaptation to an extreme habitat (vertical shale cliffs in Nuevo León, Mexico).     

Antigenic patterns of seed proteins in Opuntioideae (Cactaceae)

The antigenic patterns recognised by Western blotting in seed proteins of species of Opuntioideae (Cactaceae) were analysed in an attempt to evaluate their usefulness in systematics. Total protein profiles were also analysed by SDS–PAGE. The resulting similarity and distance matrices were further used to carry out Cluster Analysis (UPGMA) and Principal Coordinates Analysis. Populations of Opuntia cardiosperma were found to exhibit a prominent morphological uniformity, a unique electrophoretic pattern and a uniform antigenic pattern. The latter was obtained using anti-O. cardiosperma as antiserum. Results from the qualitative and quantitative interspecific analyses of antigenic profiles helped to characterise all the species studied. Tephrocactus articulatus and Cylindropuntia imbricata evidenced lower affinity with O. cardiosperma than the species of Opuntia s.s. Our results demonstrate that in Cactaceae, Western blotting analysis broadens the usefulness range of immunological techniques at the specific level and complements the information collected from electrophoretic profiles.     

Epidermal and hypodermal characteristics in North American Cactoideae (Cactaceae)

Dermal and hypodermal anatomical features of 70 species representing 21 genera of North American Cactoideae were studied. Results show that all species examined have parallelocytic stomata and anticlinal wall surface varies from straight to undulate. Cuticle thickness is mostly narrow (1–10 μm) contrary to the general opinion that cuticle is thick in most cacti; however, few species such as Ariocarpus fissuratus and several species of Pachycereus show a distinctive thick cuticle. More than 80% of the species studied have a single-layered epidermis. Papillae occur in eight species belonging to four genera. Notable papillae are a feature shared by all members of Peniocereus subg. Peniocereus. Other species show a bullate surface produced by irregular patches of secondary epidermal cell divisions. Commonly, the hypodermis is composed of more than two cell layers with distinctive collenchymatous walls as reported in many South American species. Silica bodies, prismatic crystals, druses, sphaerocrystals, and tannins are the most common cellular inclusions that distinguish several genera and appear to have taxonomic value. However, a more thorough search in species of Cephalocereus, Coryphantha, Echinocereus, Mammillaria, Neobuxbaumia, Pilosocereus, and Turbinicarpus is needed to support the previous assertion. Electronic Publication     

Orientation and its consequences for Copiapoa (Cactaceae) in the Atacama Desert

Summary Three species of the barrel cactus Copiapoa (C. cinerea, C. columna-alba, C. haseltoniana) were investigated in their native habitats along the cool, arid coastal regions of the Atacama Desert in northern Chile. All species orient towards the north with a high degree of precision. Two consequences of adaptive value result from this northerly orientation. First, tissue temperatures of the meristematic and floral regions on the tip of the cactus receive high solar radiation loads which result in high temperatures (30°–40°C) relative to air temperatures (15°–20°) during winter and spring months when adequate soil moisture for growth is available. Second, absorption of solar radiation by the sides of the cactus is minimized, which reduces both the potential detrimental effects of light and heat load on the cactus and probably balances daily quanta absorbed for photosynthesis with nighttime CO₂ uptake rates during drought stress periods.     

Vivipary and offspring survival in the epiphytic cactus Epiphyllum phyllanthus (Cactaceae)

Vivipary, the germination of seeds before they are shed from the parent plant, is a rare event in angiosperms involving complex ecophysiological processes. Pseudovivipary and cryptovivipary occur in approximately 30 (2%) species of the cactus family. A remarkable case of vivipary in Epiphyllum phyllanthus is described here. Information is provided regarding the biology of viviparous fruits, morphology, mortality, survival rates of viviparous offspring, and some eco-evolutionary implications of this reproductive strategy in the Cactaceae. This epiphytic cactus has no host-specific relationship. A low proportion (33.3%) of individuals produced viviparous fruits. Seed number/fruit varied from 197 to 230 with percentage of viviparous germination from 97.5% to 99%. The viviparous seedlings exhibited normal development and were no different from non-viviparous offspring. Transplanting experiments showed that the first week is critical for seedling establishment, and high mortality occurred in the three treatments used: 69% on the phorophyte surface, 58.6% on the ground, and 44.8% under controlled conditions. The number of survivors gradually stabilized, and the contribution to establishment was comparable in each of the treatments after the acclimation phase. It is suggested that vivipary is associated with thermoregulation, parental care, conspecific nursing, and rapid seedling establishment. Germination is not a limiting factor in the perpetuation of this viviparous species, but seedling establishment is. In viviparous individuals of E. phyllanthus, seedling mortality during establishment rather than failure to germinate within the fruit is a limiting factor affecting local population density. Overall, viviparity is an intrinsic, genetic event involving high metabolic costs favouring germination and dispersal of the fittest offspring regardless of substrate and environmental conditions.     

Radioimmunoassays of 3,4,5-trimethoxyphenethylamine (mescaline) and 2,5-dimethoxy-4-methylphenyl-isopropylamine(DOM)

Mescaline (3,4,5-trimethoxyphenethylamine) and N-succinylmescaline were coupled to human serum albumin with carbodiimide. DOM (2,5-dimethoxy-4-methylphenylisopropylamine) was linked to human serum albumin by reaction with glutaraldehyde. These conjugates were used for immunization of rabbits. Derivatives of mescaline [N-(3′,4′,5′-trimethoxyphenethyl)-4-hydroxyphenylacetamide] and of DOM [N-(2′,5′-dimethoxy-4′-methylphenylisopropyl)-4-hydroxyphenylacetamide], which could be iodinated to high specific activity, were synthesized. The antibodies bound specifically to these iodinated compounds. In competition experiments with a mescaline antiserum, 6 pmoles of mescaline inhibited 50%; with a DOM antiserum, 50% inhibition was observed with 118 pmoles of DOM. Thus, 100 pg of mescaline and 2 ng of DOM can be detected. The specificity of both antibodies is such that structurally related molecules, such as p-methoxy-phenethylamine or 3-methoxytyramine, are several orders of magnitude less effective as inhibitors than the parent molecules. With the use of antimescaline, it was determined that mescaline administered intravenously to rabbits disappeared rapidly from the circulation. The acid was identified as the major metabolite in the serum and urine of these animals.