Distribution and Chemical Composition of Lignin in Secondary Xylem of Cactaceae

Cactaceae family has heterogeneity in the accumulation of lignocellulose due to the diversity of shapes and anatomy of the wood. Most studies focus on fibrous and dimorphic species; but the non-fibrous species are poorly studied. The aims of this work were to analyze the syringyl/guaiacyl ratio of lignin and its distribution in secondary xylem, especially in non-fibrous species. The syringyl/guaiacyl (S/G) ratio was quantified from 34 species of cacti by nitrobenzene oxidation of free-extractive wood. The distribution of lignocellulose in wood sections stained with safranin O/fast green was determined with epifluorescence microscopy. The S/G ratio was heterogeneous; most of the non-fibrous species had a higher percentage of syringyl, while the fibrous ones accumulate guaiacyl. Fluorescence emission showed that vessel elements and wide-band tracheids had similar tonalities. It is hypothesized that the presence of a higher percentage of syringyl in most cacti is part of the defense mechanism against pathogens, which together with the succulence of the stem represent adaptations that contribute to survival in their hostile environments.     

Ontogenetic mechanisms and the evolution of Cactaceae

Abstract

The ancestors of cacti were leafy trees that had hard, woody trunks. The development of the cactus body is controlled by ontogenetic mechanisms that have evolved, and now they produce a body that is leafless, succulent and has a photosynthetic cortex. Specific changes include: bark formation is postponed and the epidermis and stomata function for many years; the outer cortex is a palisade cortex with intercellular spaces; there are cortical bundles that resemble leaf veins but which have secondary xylem and phloem. Wood development has changed dramatically such that water storage is maximized (increased ray parenchyma) and danger of water stress is minimized (increased paratracheal parenchyma, loss of fibers). Several genera have polymorphic wood: the plants produce one type of wood for several years, then later they produce a different type. It is possible that the extensive evolutionary changes have resulted from mutations in the controller regions of genes, not in the structural regions.     

Novel seed coat lignins in the Cactaceae: structure,distribution and implications for the evolution of lignin diversity

We have recently described a hitherto unsuspected catechyl lignin polymer (C‐lignin) in the seed coats of Vanilla orchid and in cacti of one genus, Melocactus (Chen et al., Proc. Natl. Acad. Sci. USA. 2012, 109, 1772‐1777.). We have now determined the lignin types in the seed coats of 130 different cactus species. Lignin in the vegetative tissues of cacti is of the normal guaiacyl/syringyl (G/S) type, but members of most genera within the subfamily Cactoidae possess seed coat lignin of the novel C‐type only, which we show is a homopolymer formed by endwise β–O–4‐coupling of caffeyl alcohol monomers onto the growing polymer resulting in benzodioxane units. However, the species examined within the genera Coryphantha, Cumarinia, Escobaria and Mammillaria (Cactoideae) mostly had normal G/S lignin in their seeds, as did all six species in the subfamily Opuntioidae that were examined. Seed coat lignin composition is still evolving in the Cactaceae, as seeds of one Mammillaria species (M. lasiacantha) possess only C‐lignin, three Escobaria species (E. dasyacantha, E. lloydii and E. zilziana) contain an unusual lignin composed of 5‐hydroxyguaiacyl units, the first report of such a polymer that occurs naturally in plants, and seeds of some species contain no lignin at all. We discuss the implications of these findings for the mechanisms that underlie the biosynthesis of these newly discovered lignin types.     

Effects of light and temperature on seed germination of cacti of Brazilian ecosystems

Environmental factors are used by plants as spatio‐temporal indicators of favorable conditions for seed germination. Thus, the objective of this study was to determine the effect of light and temperature on seed germination of 30 taxa of Cactaceae occurring in northeastern Brazil and to evaluate whether fluctuations in temperature are capable of altering light sensitivity. The seeds were tested for germination under two light conditions (12 h photoperiod and continuous darkness) and 10 temperature treatments: eight constant temperatures (10, 15, 20, 25, 30, 35, 40 and 45°C) and two alternating temperatures (30/20°C and 35/25°C). The species studied showed two photoblastic responses. All cacti from the Cactoideae subfamily (22 taxa) were classified as positive photoblastic (i.e., no germination in darkness), regardless of the temperature treatment used. Likewise, temperature fluctuation did not alter the seed sensitivity to light. On the other hand, the species of the Opuntioideae (five taxa) and Pereskioideae (three taxa) subfamilies are indifferent to light (i.e., germinated both in the presence and absence of light). The cacti from the areas of Caatinga and Cerrado showed an optimal germination temperature of 30°C, while the species from Atlantic Forest and Restinga areas showed an optimal germination temperature of 25°C.     

Complex biomineralization pattern in cactaceae

The infrared spectroscopic investigation of biominerals isolated from different Cactaceae species belonging to the Opuntioideae subfamily shows the presence of a very complex mineral composition, including whewellite (monohydrated calcium oxalate), opal (SiO2) and calcite (CaCO3). This is the first report on the presence of a calcium carbonate in these types of plants.     

Long-term population studies and the development of an integrated management programme for control of Opuntia stricta in Kruger National Park, South Africa

1. A cactus, Opuntia stricta , has invaded almost 16 000 ha of conserved, natural habitatand has become a major weed problem in Kruger National Park (KNP), South Africa.
2. The main objectives in the control of O. stricta are to reduce the density of the weed and to curb long-range dispersal of seeds by preventing young plants from reaching the size (28 cladodes) at which they start to produce fruits.
3. Herbicides have failed to provide satisfactory control of O. stricta because the weed infestations are replenished from seeds in the soil and from small plants that are overlooked during spraying.
4. A phycitid moth, Cactoblastis cactorum , was released in KNP during 1988 in an attempt to control O. stricta biologically.
5. Population counts of the biological control agent and of the weed over a 5-year period showed that, even though C. cactorum has not provided complete control of O. stricta in KNP, the moderate levels of larval damage have stunted the growth of O. stricta and have considerably extended the time that the young plants take to reach sexual maturity.
6. Comparisons of modelled (i.e. with no C. cactorum ) and actual populations of O. stricta showed that C. cactorum is making a substantial contribution to the control of O. stricta in residual infestations of the weed that have been treated with herbicides.
7. The need for long-term evaluation studies in biological weed control is demon strated by the development of an integrated management programme for effective control of O. stricta .     

Reproductive biology of a rare cactus, Opuntia spinosissima (Cactaceae), in the Florida Keys: why is seed set very low?

 Opuntia spinosissima (Martyn) Mill. (Cactaceae) is an extremely rare taxon with a single remaining wild population of 13 plants located on Little Torch Key, Florida. The plants rarely set viable seeds and most ovaries abscise without initiation of fruits. Pollination and breeding system were examined in Little Torch Key and in plant accessions located at Fairchild Tropical Garden. Four pollination treatments were carried out to test for apomixis, self-pollination, outcrossing, and autogamy; a control treatment was also monitored. Additionally, pollen viability, pollinators, and seed set and viability were investigated. On selfed, crossed, and open (control) stigmas pollen grains germinated, but the tubes usually did not reach the base of the style. This suggests that O. spinosissima has pre-zygotic self-incompatibility. None of the pairwise crosses set seed, so the extant plants were apparently not inter-compatible. Out of 173 manipulated and control flowers, only one set fruit. Although this flower was outcrossed, no pollen tubes germinated on the stigma; this suggests agamospermy, a process common in the Cactaceae. In those flowers where pollen tubes did reach the ovary they failed to penetrate ovules, suggesting ovarian inhibition or that this taxon has lost the ability to be fertilized. Most field-collected seeds were viable, but there is no seedling recruitment under natural conditions, and vegetative reproduction is common. Based on these findings, I hypothesize that O. spinosissima is a sterile polyploid and that the 13 extant plants are asexually derived from a single lineage. Received: 17 April 1998 / Revision accepted: 3 July 1998     

Reproductive biology and mating system estimates of two Andean melocacti, Melocactus schatzlii and M. andinus (Cactaceae)

BACKGROUND AND AIMS: The genus Melocactus comprises 36 species of globose cacti with the most derived traits in the Cereeae tribe. It is the proper study system to examine what are the most derived reproductive strategies within that tribe. This study aims to characterize the reproductive biology and to estimate the mating system parameters of two Andean melocacti, Melocactus schatzlii and M. andinus. METHODS: The reproductive attributes of the two species were described, including floral morphology, anthesis patterns, floral rewards, floral visitors and visitation patterns. Levels of self-compatibility and autonomous self-pollination were estimated by hand-pollination experiments. Mating system estimates were obtained by conducting progeny array analyses using isozymes. KEY RESULTS: The flowers of the two species present the typical hummingbird-pollination syndrome. Despite their morphological resemblance, the two species differ in flower size, pollen and ovule production and anthesis pattern. Their main pollinator agents are hummingbirds, four species in M. schatzlii and one species in M. andinus. Both cacti are self-compatible and capable of self-pollination without the aid of pollen vectors. Population-level outcrossing rate was higher for M. schatzlii (t(m)=0.9) than for M. andinus (t(m)=0.4). At the family level, outcrossing rates for most mothers of M. schatzlii were higher (t(m)>0.8) than for M. andinus (t(m)<0.5). CONCLUSIONS: Although the two cacti are capable of selfing, M. schatzlii is a predominantly outcrossing species, while M. andinus behaves as a mixed-mating cactus. Hummingbirds are the only pollinators responsible for outcrossing and gene flow events in these species. In their absence, both melocacti set seeds by selfing. Based on its low population size, restricted distribution in Venezuela, low rates of floral visits, and high levels of inbreeding, M. andinus is considered to be an endangered species deserving further study to define its conservation status.