Irrigation magnifies CAM-photosynthesis in Opuntia basilaris (Cactaceae)

Summary Measurements of acid metabolism and gas exchange were carried out four times during a year to assess the relative importance of temperature and the accompanying seasonal change to the carbon metabolism of Opuntia basilaris Engelm. & Bigel. plants growing in situ under irrigated and natural (control) conditions. Our experiments showed that this cactus did not change its pattern of carbon assimilation when continuously irrigated under field conditions. Non-irrigated cacti had maximum acid accumulation after periods of precipitation. Maximum acid accumulation in irrigated cacti occurred when there was a large difference between day/night temperatures (i.e., 16°C), and when nighttime temperatures were moderate (14C). Irrigated cacti had greater duration of stomatal opening and lower resistance to ¹⁴CO₂ uptake. When temperatures were low, daytime stomatal resistance to ¹⁴CO₂ uptake decreased (to 20–40 s cm^-1), but never to the level of the nocturnal resistances (5–10 s cm^-1). During periods of drought, nonirrigated cycti changed to a pattern in which organic acids fluctuated. Irrigated cacti continued to have ¹⁴CO₂ uptake when nighttime temperatures were as high as 33°C. ¹³C/¹²C isotope composition ratios, determined after two years of irrigation, were near -12 in irrigated and non-irrigated plants. Therefore, under conditions of continual irrigation, seasonal and temperature changes affected the degree of dark CO₂ fixation and acid metabolism, but these cacti did not change from CAM to CO₂ fixation in the light.Abbreviations C₃ reductive pentose phosphate cycle - C₄ dicarboxylic acid cacle - CAE carbon assimilation efficiency - CAM Crassulacean acid metabolism - THO tritiated water - T/P transpiration ratio - vpd vapor pressure deficit - water potential This study was supported in part by National Science Foundation grant OIP 74-15673. Under the auspice of this grant a cooperative research project was carried out between the Australian National University, Canberra, and the Philip L. Boyd Deep Canyon Desert Research Center, University of California, Riverside. The studies involved a comparison of the photosynthetic pathways employed by succulents during growth in their native environment (the Sonoran Desert of southeastern California) and in a favorable introduced environment (Queensland and New South Wales, Australia). Studies carried out in Australia are under the direction of Dr. C.B. Osmond     

Spine influences on PAR interception, stem temperature, and nocturnal acid accumulation by cacti

Abstract. The influence of spines on nocturnal acid accumulation was examined for two Crassulacean acid metabolism plants, the barrel cactus Ferocactus acanthodes (Lemaire) Britton & Rose var. lecontei (Engelm.) Lindsay and the cylindropuntia Opuntia bigelovii Engelm. var. bigelovii. The ambient photo-synthetically active radiation (PAR) required for 90% saturation of nocturnal acid accumulation by O. bigelovii was 23 mol m^?2 d^?1 for stems that were 32% shaded by spines and 16 mol m^?2 d^?1 when the spines were removed. For F. acanthodes, 90% saturation occurred for an ambient PAR of 45 mol m^?2 d^?1 where the stem was 78% shaded by spines and 21 mol m^?2 d^?1 when the spines were removed. For comparison, the ambient PAR averaged 21 mol m^?2 d^?1 at various times of the year and stem positions for O. bigelovii fully exposed to solar irradiation. Although spines can reflect some PAR toward the stem surface, their presence decreases the PAR incident on the stem surface, thereby decreasing the nocturnal acid accumulation. For O. bigelovii, periodically removing the spines led to a 60% greater increase in stem volume over 2^1/2 years. The effect of variation in spine-shading on stem surface temperature was determined for F. acanthodes using computer simulations and an energy budget model. A substantial reduction in daytime stem temperatures and a smaller increase in night-time temperatures as the spine-shading increased had very little influence on nocturnal acid accumulation for both winter and summer days. Thus, the main effect of spines on the metabolism of cacti is apparently to reduce the PAR incident on the stem surface and thereby to reduce productivity.     

Relationships between Stomatal Behavior and Internal Carbon Dioxide Concentration in Crassulacean Acid Metabolism Plants

Measurements of internal gas phase CO₂ concentration, stomatal resistance, and acid content were made in Crassulacean acid metabolism plants growing under natural conditions. High CO₂ concentrations, sometimes in excess of 2%, were observed during the day in a range of taxonomically widely separated plants (Opuntia ficus-indica L., Opuntia basilaris Engelm. and Bigel., Agave desertii Engelm., Yucca schidigera Roezl. ex Ortiges, Ananas comosus [L.] Merr., Aloe vera L., Cattleya sp. and Phalanopsis sp.) and below ambient air concentrations were observed at night.     

THE DEVELOPMENTAL ANATOMY OF OPUNTIA BASILARIS. I. EMBRYO,ROOT, AND TRANSITION ZONE

The large seeds of Opuntia basilaris Engelm. & Bigel. show an unusually high percentage of germination, followed by a rapid development of the seedling during the first 30 days of growth. The primary root has six xylem arms alternating with six phloem poles around a large central pith. Development of metaxylem opposite each of the primary phloem poles results in the formation of eight collateral bundles. Secondary and tertiary roots have four xylem and phloem poles with xylem developing to the center of the stele. The transition zone is characterized by a gradual disappearance of all but two of the primary xylem arms of the root. Metaxylem development in the central portion of the transition zone interconnects the protoxylem poles forming a primary xylem cylinder around the central pith. The collateral bundles pass through the transition zone essentially without change.     

Apical meristem organization and lack of establishment of the quiescent center in Cactaceae roots with determinate growth

Some species of Cactaceae from the Sonoran Desert are characterized by a determinate growth pattern of the primary root, which is important for rapid lateral-root formation and seedling establishment. An analysis of the determinate root growth can be helpful for understanding the mechanism of meristem maintenance in plants in general. Stenocereus gummosus (Engelm.) Gibson & Horak and Pachycereus pringlei (S. Watson) Britton & Rose are characterized by an open type of root apical meristem. Immunohistochemical analysis of 5-bromo-2-deoxyuridine incorporation into S. gummosus showed that the percentage of cells passing through the S-phase in a 24-h period is the same within the zone where a population of relatively slowly proliferating cells could be established and above this zone in the meristem. This indicated the absence of the quiescent center (QC) in S. gummosus. During the second and the third days of growth, in the distal meristem portion of P. pringlei roots, a compact group of cells that had a cell cycle longer than in the proximal meristem was found, indicating the presence of the QC. However, later in development, the QC could not be detected in this species. These data suggest that during post-germination the absence of the establishment of the QC within the apical meristem and limited proliferative activity of initial cells are the main components of a determinate developmental program and that establishment of the QC is required for maintenance of the meristem and indeterminate root growth in plants.Abbreviations QC quiescent center - RCP root cap-protoderm - BrdU 5-bromo-2-deoxyuridine - FITC fluorescein isothiocyanate - DAPI 4,6-diamidino-2-phenylindole     

DEVELOPMENTAL ANATOMY OF EPIDERMAL AND MESOPHYLL CHLOROPLASTS IN OPUNTIA BASILARIS LEAVES

Developmental studies of Opuntia basilaris Engelm. & Bigel. leaves revealed the presence of three morphologically distinct types of plastids. All epidermal cells examined contained chloroplasts. After 13 days of growth in the dark the plastids of epidermal and mesophyll cells were characterized by the presence of a prolamellar body and fibril inclusions. Epidermal plastids which developed under light conditions contained large stromacentres and a limited grana-fret membrane system. Guard cell plastids developed under similar conditions had a much poorer developed grana system with fibril inclusions apparent only during the developmental stages. At maturity these plastids appeared swollen or dilated. Mesophyll plastids had fibril inclusions during all stages of development and at maturity contained a very extensive grana-fret membrane system. Microbodies were found in association with the mesophyll plastids. Starch accumulation was common in subsidiary cell and guard cell plastids.     

Influence of vesicular-arbuscular mycorrhizae on biomass production by the cactus Pachycereus pecten-aboriginum

This study reports the effect of vesicular-arbuscular mycorrhizal (VAM) fungi on dry matter production by Pachycereus pecten-aboriginum (Engelm.) Britt & Rose, an arborescent cactus of arid and tropical dry forest in Mexico. Seedlings in the presence or absence of VAM fungi were grown in soil between two plates of glass (20 × 30 cm) for 8 months inside growth chambers (30/25° C, 13/11 h day/night and a light intensity of 400 mol m^-2 s^-1). VAM seedlings had significantly (P<0.01) higher dry matter production (0.418 versus 0.169 g), root/shoot ratios (0.26 versus 0.14) and specific root length (0.65 versus 1.41 mm mg^-1) than non-VAM seedlings, suggesting a more efficient exploitation of soil resources by the VAM cacti. The data point to a role for VAM fungi in the establishment, growth, water relations and nutrition of cacti in the arid tropics.     

ORIENTATION,PAR INTERCEPTION,AND NOCTURNAL ACIDITY INCREASES FOR TERMINAL CLADODES OF A WIDELY CULTIVATED CACTUS,OPUNTIA FICUS-INDICA

Terminal vertical cladodes (flattened stems) of Opuntia ficus-indica growing in widely separated locations were nonrandomly oriented. On plantations at 33°S latitude in Chile individual cladodes tended to orient in the same direction as the planted cladodes on which they developed. However, after 2 years unshaded new cladodes tended to face east-west. Terminal cladodes also tended to face east-west for irrigated O. ficus-indica in California (at 34°N) and in Israel (32 to 33°N), but cladodes developing in the winter tended to face north-south. Except for the residual effect of initial planting direction, the observed patterns tended to maximize the interception of photosynthetically active radiation (PAR). Specifically, east-west cladode orientation would maximize PAR interception, except for cladodes developing near the winter solstice at latitudes more than 27° from the equator. Nocturnal acidity increases and hence productivity would generally be light-limited, since the nocturnal increase in acidity was 90% saturated for a total daytime PAR of 24 mol m^–2 day^–1 and the PAR received on vertical surfaces is usually less than this. Topographical features can modify the orientation patterns, since at a site where PAR was considerably blocked by surrounding mountains the maximal nocturnal acidity increases and peak in cladode orientation occurred 20° from facing east–west. Laboratory studies showed that developing cladodes oriented toward a horizontal light and were rotated an average of 16° in a direction that increased PAR interception compared to the cladodes on which they developed. Such phototropic responses, the higher productivity of favorably oriented cladodes, and the tendency to orient similarly to the underlying cladode presumably accounts for the overall orientation patterns observed, where up to four times more cladodes may face in a particular direction than at right angles to it.     

Phosphoenolpyruvate carboxylase from spinach leaf tissue: inhibition by sulfite ion

Respiration and gas exchange in the light were studied manometrically with tissue slices from stem material of Opuntia basilaris Engelm. and Bigel. Dark respiration rates were greater in young stems than in mature stems. The timing of the experiment in the day/night cycle influences the magnitude and pattern of respiration and gas exchange in the light. Net dark respiration has a temperature optimum between 35 and 40 C, and is maintained at 60% of the control rate in tissue equilibrated with experimental osmotic potentials of −25 bars. Net gas exchange in the light is regulated by the titratable acidity of the tissue and by the tissue temperature. Increased rates of net CO₂ evolution and net O₂ consumption occur in the light with high levels of titratable acidity and high temperatures. An efflux of CO₂ and influx of O₂ occur following light/dark transitions. These patterns are reversed following dark/light transitions. Similar results were demonstrated at 15, 25, and 35 C, and are interpreted as a mechanism of adaptation to desert environments.     

Seasonal Patterns of Acid Metabolism and Gas Exchange in Opuntia basilaris

Acid metabolism and gas exchange studies were conducted in situ on the cactus Opuntia basilaris Engelm. and Bigel. A pattern of significant seasonal variation was evident. The pattern was controlled by rainfall, which significantly influenced plant water potentials, total gas transfer resistances, and nocturnal organic acid synthesis. In winter and early spring, when plant water stress was mild, stomatal and mesophyll resistances remained low, permitting enhanced nocturnal assimilation of ¹⁴CO₂. The day/night accumulation of acidity was large during these seasons. In summer and fall, plant water stress was moderate, although soil water stress was severe. The nocturnal assimilation of ¹⁴CO₂ was very low during these seasons, even in stems with open stomata, indicating large mesophyll resistances restricting exogenous gas incorporation. The day/night accumulation of acidity was reduced, and a low level of acid metabolism persisted throughout this period. The rapid response to a midsummer rainfall emphasizes the importance of plant water potential as a parameter controlling over-all metabolic activity. The seasonal variations of acid metabolism and gas exchange significantly influenced the efficiency of water use and carbon dioxide assimilation. Periods of maximal efficiency followed rainfall throughout the course of the year.     

INTERACTIVE EFFECTS OF PAR AND UV‐B RADIATION ON PSII ELECTRON TRANSPORT IN THE MARINE ALGA DUNALIELLA TERTIOLECTA (CHLOROPHYCEAE)1

To better understand the interactions between PAR and UV‐B radiation in microalgae, the marine chlorophyte alga Dunaliella tertiolecta was subjected to a UV‐B flux of 4.1 W·m ^{? 2} (unweighted) with varying PAR fluxes. Rate constants for damage and repair processes during UV‐B exposure increased with PAR flux. However, recovery after UV‐B exposure increased with PAR up to 300 μmol quanta·m ^{? 2}·s ^{? 1} 1 Received 17 September 2002. Accepted 19 February 2003. , beyond which photoinhibition of PSII electron transport was found to decrease recovery rates. In the absence of PAR during the post UV‐B exposure period, no recovery was seen, indicating that perhaps the lack of light available for photosynthesis depresses repair either directly or indirectly by affecting ATP synthesis. Possible mechanisms for the observed interactions between PAR and UV‐B exposure are discussed.     

Carbon assimilation patterns and growth of the introduced CAM plant Opuntia inermis in Eastern Australia

Summary The daily course of CO₂ and H₂O exchange in cladodes of Opuntia inermis was studied at four sites in Eastern Australia. On most occasions cladode water contents were high and nocturnal stomatal opening resulted in substantial uptake of CO₂ and synthesis of about 130 equiv cm^-2 of malic acid during the night. Under water stress nocturnal stomatal opening was confined to the latter part of the night and acid synthesis was reduced to about 40 equiv cm^-2. Night temperature had little effect on acid synthesis, which responded primarily to rainfall and changed from the stressed condition within 2–3 days in irrigation experiments. On many occasions following summer rainfall stomata opened for 4 h in the late afternoon permitting net CO₂ fixation which may contribute about 25% of the total carbon assimilated. This CO₂ fixation was insufficient to have a marked impact on the ₁₃C value of the Opuntia cladodes. CO₂ fixation in the light occurred in conjunction with maximum dark CO₂ fixation under mesic conditions. Dark CO₂ fixation rates were 3 to 5 times greater than those recorded in desert cacti under favorable conditions. Relative growth rates calculated on the basic of CO₂ exchange correspond to measured relative growth rates of 0.05 g g^-1 dry wt day^-1 which prevailed for 60–90 days in summer. The capacity for very active CO₂ fixation in the dark and light following summer rainfall and the capacity to persist at low levels of metabolic activity through summer drought are discussed in relation to the success of this introduced species in this habitat.     

The Echinocereus enneacanthus-dubius-stramineus complex (Cactaceae)

A taxonomic treatment of three species of Texas desert and desert-margin cacti,Echinocereus enneacanthus Engelm.,E. dubius (Engelm.) Rümpler, andE. stramineus (Engelm.) Rümpler, is presented. Field studies show thatE. enneacanthus is a grassland and desert-margin species rarely found above altitudes of 2000 ft in Texas, whileE. dubius andE. stramineus are desert-margin and desert species not found below 2000 ft. Morphological studies make possible the identification of these species on the basis of the shape, size, and arrangement of the spines. The epidermal characteristics are discussed, and the possibility of the use of the epidermal cell patterns as identifying characters is postulated. The speciesE. enneacanthus is divided into three forms based on spine characters and habitat.     

Efficient transformation of potato (Solanum tuberosum L.) using a binary vector in Agrobacterium rhizogenes

Summary We transformed three potato (Solanum tuberosum L.) genotypes by using A. rhizogenes or a mixture of A. rhizogenes and A. tumefaciens. Inoculations of potato stem segments were performed with Agrobacterium rhizogenes AM8703 containing two independent plasmids: the wild-type Ri-plasmid, pRI1855, and the binary vector plasmid, pBI121. In mixed inoculation experiments, Agrobacterium rhizogenes LBA1334 (pRI1855) and Agrobacterium tumefaciens AM8706 containing the disarmed Ti-plasmid (pAL4404) and the binary vector plasmid (pBI121) were mixed in a 11 ratio. The T-DNA of the binary vector plasmid pBI121 contained two marker genes encoding neomycin phosphotransferase, which confers resistance to kanamycin, and -glucuronidase. Both transformation procedures gave rise to hairy roots on potato stem segments within 2 weeks. With both procedures it was possible to obtain transformed hairy roots, able to grow on kanamycin and possessing -glucuronidase activity, without selection pressure. The efficiency of the A. rhizogenes AM8703 transformation, however, was much higher than that of the mixed transformation. Up to 60% of the hairy roots resulting from the former transformation method were kanamycin resistant and possessed -glucuronidase activity. There was no correlation between the height of the kanamycin resistance and that of the -glucuronidase activity in a root clone. Hairy roots obtained from a diploid potato genotype turned out to be diploid in 80% of the cases. Transformed potato plants were recovered from Agrobacterium rhizogenes AM8703-induced hairy roots.     

BIOMECHANICS OF CLADODES AND CLADODE-CLADODE JUNCTIONS FOR OPUNTIA FICUS-INDICA (CACTACEAE)

Shoots of Opuntia ficus-indica (L.) Miller consist of a sequence of flattened stem segments (cladodes) in contact over only a small portion of their periphery. The maximum angular deflection for an upper terminal cladode under a weight equal to its own fresh mass applied perpendicular to its face increased from 5° to 9° as cladode length increased from 20 cm to 60 cm, consistent with an increase in mass proportional to length^2.83. Just over half of the angular deflection of an upper cladode represented flexure of the cladode-cladode junction; the angular deflections averaged fourfold more for mass loadings perpendicular to the cladode face compared with those parallel to it. Compared with such static loading by mass, dynamic loading by wind for a 31-cm-long cladode led to a maximum angular deflection of only 0.13° at a wind speed of 1 m sec⁻¹ and 2.3° at 10 m sec⁻¹. Drought caused the angular deflections to increase 9% for 21-cm-long cladodes over a 90-day period and to decrease 45% for 44-cm-long cladodes. Increases in stem temperature from 0 C to 20 C increased angular deflections of a 27-cm-long cladode about 10%, with little further increase up to 50 C. Even though the cladodes were thin compared with stems of many perennials and the cladode-cladode junction comprised only about 3.5% of their peripheral area, the shoots of O. ficus-indica proved to be quite rigid, as angular deflections of cladodes were only slightly influenced by temperature and wind, were not markedly enhanced by drought, and were less than 10° under loading by a cladode's mass.     

Growth of and quassin accumulation by cultures of Quassia amara

Callus and suspension cultures were established from Quassia amara, a member of the Simaroubaceae. Analysis of the tissue culture showed that quassin was present in both callus and suspension cultures. The effect of variation in auxin and cytokinins on both callus growth and the presence of quassin was examined. The suspension culture was grown in a 7 liter bioreactor when good yields of quassin were achieved.Abbreviations IAA indole-3-acetic acid - IBA indolebutyric acid - 2,4-D 2,4-dichlorophenoxyacetic acid - NAA naphthaleneacetic acid - 6BA 6-benzyladenine - IpA N⁶ (-isopentenyl) adenine - IpAR N⁶ ( isopentenyl) adenine riboside - td doubling time     

Specificity of siderophore-mediated transport of iron in rhizobia

The trihydroxamate siderophore, hydroxamate K, has been purified from culture filtrates of iron-deficient Rhizobium leguminosarum biovar viciae MNF710. The iron complex has a molecular weight of 828 and an absorption maximum at 443 nm (_M=1510). ⁵⁵Fe complexed to purified hydroxamate K was taken up by MNF710, its hydroxamate-negative mutant MNF7102 and Rhizobium leguminosarum biovar trifolii WU95 via an iron-regulated transport system, but Rhizobium meliloti U45 failed to take up the iron-siderophore complex under any conditions. A similar pattern of iron uptake was observed with ferrioxamine B. MNF710, MNF7102, U45 and WU95 all transported ⁵⁵Fe-ferrichrome but only the first three strains took up ⁵⁵Fe-ferrichrome A. All these ⁵⁵Fe-trihydroxamate uptake systems were ironregulated in MNF710, MNF7102 and WU95. In contrast, uptake of ⁵⁵Fe-rhodotorulate, a dihydroxamate, was essentially constitutive in all four organisms. Similarly, uptake of ⁵⁵Fe-citrate and ⁵⁵Fe-nitrilotriacetic acid was constitutive. None of the strains took up ⁵⁵Fe complexed with enterobactin or with pyoverdins from Pseudomonas aeruginosa ATCC15692 (PAO1) and Pseudomonas fluorescens ATCC17400.     

Past climate changes and ecophysiological responses recorded in the isotope ratios of saguaro cactus spines

The stable isotope composition of spines produced serially from the apex of columnar cacti has the potential to be used as a record of changes in climate and physiology. To investigate this potential, we measured the δ¹⁸O, δ¹³C and F¹⁴C values of spines from a long-lived columnar cactus, saguaro (Carnegiea gigantea). To determine plant age, we collected spines at 11 different heights along one rib from the stem apex (3.77 m height) to the base of a naturally occurring saguaro. Fractions of modern carbon (F¹⁴C) ranged from 0.9679 to 1.5537, which is consistent with ages between 1950 and 2004. We observed a very strong positive correlation (r = 0.997) between the F¹⁴C age of spines and the age of spines determined from direct and repeated height measurements taken on this individual over the past 37 years. A series of 96 spines collected from this individual had δ¹⁸O values ranging from 38‰ to 50‰ [Vienna standard mean ocean water (VSMOW)] and δ¹³C values from ?11.5‰ to ?8.5‰ [Vienna Peedee belemnite (VPDB)]. The δ¹⁸O and δ¹³C values of spines were positively correlated (r = 0.45, P < 0.0001) and showed near-annual oscillations over the ～15-year record. This pattern suggests that seasonal periods of reduced evaporative demand or greater precipitation input may correspond to increased daytime CO₂ uptake. The lowest δ¹⁸O and δ¹³C values of spines observed occurred during the 1983 and 1993 El Niño years, suggesting that the stable isotope composition recorded in spine tissue may serve as a proxy for these climate events. We compared empirical models and data from potted experimental cacti to validate these observations and test our hypotheses. The isotopic records presented here are the first ever reported from a chronosequence of cactus spines and demonstrate that tissues of columnar cacti, and potentially other long-lived succulents, may contain a record of past physiological and climatic variation.     

Leaf photosynthetic characteristics of seedlings of actinorhizal Alnus spp. and Elaeagnus spp.

Single leaf photosynthetic characteristics of Alnus glutinosa, A. incana, A. rubra, Elaeagnus angustifolia, and E. umbellata seedlings conditioned to ambient sunlight in a glasshouse were assessed. Light saturation occurred between 930 and 1400 mol m^-2s^-1 PAR for all species. Maximum rates of net photosynthesis (Pn) measured at 25°C ranged from 12.8 to 17.3 mol CO₂m^-2s^-1 and rates of dark respiration ranged from 0.74 to 0.95 mol CO₂m^-2s^-1. These values of leaf photosynthetic variables are typical of early to midsuccessional species. The rate of Pn measured at optimal temperature (20°C) and 530mol m^-2s^-1 PAR was significantly (p<0.01) correlated with leaf nitrogen concentration (r=0.69) and negatively correlated with the mean area of a leaf (r=–0.64). We suggest that the high leaf nitrogen concentration and rate of Pn observed for Elaeagnus umbellata and to a lesser degree for E. angustifolia are genetic adaptations related to their crown architecture.Abbreviations Pn net photosynthesis     

Stem tilting in the inter‐tropical cactus Echinocactus platyacanthus: an adaptive solution to the trade‐off between radiation acquisition and temperature control

While plants require radiation for photosynthesis, radiation in warm deserts can have detrimental effects from high temperatures. This dilemma may be solved through plant morphological attributes. In cold deserts, stem tilting keeps reproductive organs warm by increasing radiation interception at the cost of decreased annual light interception. Conversely, little is known about stem tilting in warm deserts. We hypothesised that stem tilting in Echinocactus platyacanthus prevents high temperatures near the apex, where reproduction occurs. The study was conducted in the warm, inter‐tropical portion of the Chihuahuan Desert, Mexico. We found that cacti preferentially tilted towards the south, which reduced temperatures of reproductive organs during the hot season, but increased total annual near‐apex PAR interception. Tilting also maximised reproduction, a likely consequence of temperature control but perhaps also of the difficulty in translocating photosynthates in cacti; therefore, annual energy acquisition near floral meristems may be largely allocated to reproduction. Unlike plants of higher latitudes, in inter‐tropical deserts sunlight at noon comes either from the north or the south, depending on the season, and thus stem tilting may more strongly affect total annual radiation received in different portions of the stem. Inter‐tropical cacti can synchronise reproduction with irradiance peaks if flowering occurs in a specific (north or south) portion of the stem; also, they effectively solve the conflict between maximising annual PAR interception and minimising temperature at the hottest time of day. Notably, the two inter‐tropical cacti in which stem tilting has been studied successfully solve this conflict.